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B. S. Doane College, 1907 
A. M. University of Nebraska, 1909 


Submitted in Partial Fulfillment of the Requirements for the 

Degree of 






191 1 





Recommendation concurred in: 

1 37(774 

Digitized by the Internet Archive 
in 2013 



Introduction 1 

Historical 4 

The type genus 6 

The family Proteccephalidae defined 10 

Definitions of genera 10 

Anatomical 13 

Location of genital organs 13 

The fifth sucker 13 

Character of data of diagnostic value 16 

External measurements 16 

Lata on internal organs 17 

Specific description 18 

Proteocephalus ^ilicollis (Rud) 19 
" dubius nov. spec. 

,T fall ax " 47 

" longicollis (Rud} 63 

" cernuae (Gmelin) La Rue 77 

" neglectus nov. spec. 88 

osculatus (Goeze) sp.inq. 93 

" pinguis nov. spec. 1C2 

" pusillus word 113 

■ macrocephalus (Creplin) 125 




nov. spec. 








nov. spec. 




nov. spec. 




(Barbieri ) 



saline nis-umblae 

fMontieelli ) 




(O.F. Mailer) 




(Batsch ) 



pent as tonus 

(Xlaptocz ) 



f ossatus 





( Gui. 3( lmeider ) 




( Grimm) 




(Klaptocz ) 




(v. Linstov/) 







(Leidy ) 










(v. Linstov/) 


nemat osomus 

(Leidy ) 



perspi cua 

(Riggenbach,1896 )- 

La Rue 
nov. spec. 



f ilaroides 

La Rue 




( Fuhrmann ) 



g andi.s 

nov. spec. 




(Barrcis ) 


Ophio taenia Ilattereri (Parcna) 331 

" Calmettei (Barrcis) 356 

" trimeresuri(Parona) 347 

" racemosa (R:.d. ) 355 

" punica ( Cholodkovsky) 365 

" lactea (Leidy) 370 

" pigmentata (v. Linstov/) 371 

Crepidbothritun Gerrardii (Baird) 373 

O&choristica cryptobothrium { v.Linstov/) La Hue 400 

•Tetracotylus c^ry uhicephalus Mont ic ell i 401 

" Diesingii " 415 

" macrocatylea " 417 

" malopterur j " 410 

Bibliography 424 

Explanation of plates 

Plates I-IX. 



(With nine plates, one-hundrea-ninefcy-f ive figures) 


Pour years ago while a student under Prof. Henry B. Ward 
at the University of ITebraska the writer "began an investigation of 
a cestode, parasitic in Amblystoma tigrinum Baird, an amphibian 
common to Nebraska. This investigation resulted in a paper (La 
Rue, 1909) in which several problems to "be investigated at an early 
date were outlined. With the problem of histogenesis of Proteoeeph - 
a lus filaroides La Rue in view a considerable amount of material 
was collected but not sufficient for the completion of that work. 
My removal from Nebraska to Illinois put an end to further collect- 
ion of the parasite, and altho several collecting trips were made 
to Nebraska and Kansas no more parasites were secured. 

Meanwhile Professor T/ard whose collection already contained 
many specimens of Proteocephalus had been adding to the number by 
collections in the field and by exchange. The writer had also made 
collections of several species* 

A preliminary study of the available material convinced 
Professor Ward that a more complete and comparative study of the 
genus was desirable and that the results of such a study would 
prove of value to helminthologist s , not only in America but also in 
Europe. Such a study seemed more desirable since but two compara- 
time studies of the genus had been made in fifteen years, the 
first by Riggenbach (1896) mostly on the species infesting fish, 


and the second by Schwarz (1908) wholly on the species infesting 
snakes and other reptiles. The fact that very little had been done 
on the genus by American investigators furnished an added incentive 
to undertake the work. 

At Professor 7/ard's suggestion and under his direction I 
undertook an investigation of the genus which would be comprehen- 
sive in scope. For the purposes of the work Professor 7/ard aug- 
mented the material at hand by securing the loan of alcoholics and 
prepared slides from several European and American investigators 
and from the collections of the Smithsonian Institution and the 
Bureau of Animal Industry at Washington. Unfortunately specimens 
of certain material could not be secured because it had been lost 
or because it could not be removed from "Che collections. 

Thanhs to Prof. Fritz Zschokke, Prof. G. Parona, Prof. Max 
Braun, Prof. K. M. Levander, Prof. Anton Collin, Prof. A. E. 
Shipley, Prof. F. 3. Monticelli and Prof. 0. Fuhrmann who at 
Professor 7/ard's request sent specimens of European species and 
thanks to Dr. C. W. Stiles, Dr. B. H. Ransom, Prof. E. Linton, Dr. 
A. J. Smith and Professor T. L. Hank ins on who upon request sent 
specimens of American species I have had a large amount of material 
for comparative study. 

I wish also to thank ttr. E. G. Davis of Lincoln, Nebraska, 
and Mr. Hermann Douthitt of Sulphur, Oklahoma, who have furnished 
material for study. 

I desire to thank Mr. S. Fred Prince for re-drawing many 
figures fyflvviotier works and for making my pencil sketches into the 
finished drawings. 


To Professor H. B. Ward my sincerest thanks are due for the 
use of his extensive collections, for the use of his library, and 
also for securing for ray use many rare specimens and rare "books. 
His interest and co-operation have given me the needed inspiration 
to complete the v/ork. 



Rudolph! (1808-10) collected together the results of the 
systematic labors of helminthologi sts up to that time. That work 
is very complete and in it are to he found the diagnoses synonymy 
and descriptions of the species of this genus that were known at 
that time. These species are included in the genus Taenia Linneaus, 
(1758). It is to he noted that Rudolphi renamed certain of these 
species which had been previously established by other investigators, 
^ater Ru&olphi (1819) brought down to date a summary of the in- 
vestigations of preceding helminthologists. 

Some of the early investigators other than Hudolphi, who 
reported species, now included in the genus Proteocephalus , were 
Goeze, Mailer 0. F , Pallas, Gmelin, Batsch, Schrank, Zeder, Blocfa 
and Froelich. Llany of their descriptions and diagnoses were very 
short; a few only were accompanied by drawirgs. 

It is needless to say that the work of these early investi- 
gators is almost entirely concerned with the study of external 
characters , 

There were no more important works which have to do with 
this group of cestodes until the time of Dujardin (1845) and 
Diesing (1850). These investigators listed several new species and 
nev; host species. 

Investigators up to the time of Llonticelli (1891) with the 
exception of "'einland (1058) considered this group of fish cestodes 
to belong to the genus Taenia. The latter author proposed the 


name Proteocephalus which will he discussed later and the former 
proposed the name Tetracotylus ♦ 

It was von Linstow (1891) who first pointed out that the 
fish Taenia made up a closely related group within the genus Taenia . 
He made the first careful study of one of the species of this 

Mont ie ell i (1891) made a careful study of the group and 
pointed out several misinterpretations relating to the genital 
organs. He first showed "she true relationships of the organs of 
the interovarial space. His work was largely based on Taenia 
coryphicephalQL with which he compared other species. In all he 
listed 20 species belonging to, his genus Tetr acotylus . Of these 
species some were from snakes. For a more complete discussion of 
that work see the description of T etracotylus . 

Since this work of Mont i c ell i; Lonnberg (1894) proposed the 
genus I chthyotaenia and to this genus many species have been as- 
cribed. The range of hosts includes amphibia, snakes, lizards and 
all the larger groups of freshwater fish. 

In 1899 i.ionticelli recognized that Tetrabothrium Gerrardii 
Baird, a parasite of the Boidae . belonged in this family of 
cestodes but that it should be separated from the species of Proteo - 
cephalus . and for this species he proposed the genus Crepidobffthrium , 

In 1903 von Linstow proposed the genus Acanthotaenia to in- 
clude a spiney headed Proteocephalid frorr. Varanus. This genus was 
later emended by Johnston (1909) v/ho placed three ether species in 


the genus, all from Varanidae . 

I have fcund it necessary to restrict the genus Proteoce - 
phalus "by removing from it the species of Amphibian and Ophidian 
Proteocephalids. These . ake a fairly homogenous group for which 
I propose the name Ophiotaenia IIov. gen. with Ophio taenia perspicua 
Nov. spec, as the type species. A definition of this nev; genus 
appears on page . The type species is described on pp 

Discussion of name, 
"einland (1858:53) proposed the genus P r o t e o c e p h a 1 u s and 
named as its type Taenia ambigua Dujardin. Taenia f ilicollis and 
Taenia dispar were also included under this genus. 

Lflnnberg (1894:801-803) proposed the generic name 
I cht hyo taenia and listed as types of this genus; 
I. filicollis Hud. 
I. ocellata Paid. 
I. longicollis Hud. 
I. torulosa Batsch. 
I. coryphiceph(\la 
Since Ldimberg named I. filicollis first in his list that 
name is to be considered the type of the genus and has been so 
designated by Hall (1910). Thus he made I chthyo taenia a synonym 
of Proteocephalus unless _T. filicollis Hud. is generically 
different from T. ambigua (Lujardin). Railliet (1899) says: 


"Comme cette espece ( Taenia ambigfla Du j . ) rentre nettement dans le 
geHQfe I cht hyo taenia il est evident que le premier nom ( Proteocephalus ) 
doit etre repris." 

The name Proteocephalus cannot he considered to he pre- 
occupied on account of a family name Proteocephala proposed by de 
Blaineville (1828) for a cestode family. The names are not identi- 
cal, consequently, not homonyms and the prior use of a name for a 
family does not invalidate its use later as a generic name. 

The validity of the names Pro teocephalus V.einland or 
Ichthyotaenia LSnnberg then depend on the species JD. ambigua 
Dujardin and T. f ilicollis Hud. The first named species is fairly 
well described, and measurements of diagnostic value were given by 
Dujardin (1845). This species^ has since been pretty well described 
and figured by G, Schneider and later by myself using Schneider's 
material agrees almost perfectly with Dujaruin's description of T\ 
ambigua . 

3ud. is not a synonym of Taenia ocellata . Hud. and conse- 
quently, of Taenia percae Uuller, as Kraemer, Riggenbach, Benedict, 
Railliet, an^ others have thought. That -belief was founded on the 
mistaken indentif ication of Xraemer (1892) 

Since Dujardin (1845) there have been but few records of 


exammationSfl Gajstcrosteus . One of these records is by Lbnnberg, 
another by Schneider. They examined Gasterosteus pungitius . whi/e 
the type host, of T. filictfllis is Gasterosteus aculeatus and so 
far as I can discover there have been almost no records of collec- 
tion of parasites from that host since Dujardir (1845). 


The diagnoses of jP. f ilicollis much resemble those of T. 
ggjblgoa , altho Dujardin evidently thought the tv/o species to he 
different for he records both. At the present time but a single 
species is well known from Gasterosteus , viz: I consider, however , 
Proteocephalus ambiguus (Duj.) is identical with P. f ilicollis (Rud) 
and hence that the genera Proteocephalus 7/eiland and Ichthyotaenia 
being based on the same species. 

I have thus far not considered the name Tetracotylus 
Llonticelli (1891) which was based upon the description of Taenia 
coryphicephala -lonticelli failed to desigante a type species. 

Brdtan (1894-1900) stated that this genus was based on T. 
coryphicephala and this action has beer considered by Hall (1910) to 
be equivalent to designation of a type species. Brown regards 
Tetracotylus a synonym of Ichthyotaenia which makes it a synonym 
of Proteocephalus provided that the type of Tetracotylus belongs 
in the same genus with Taenia f ilicollis (Rud), the type of 
Proteocephalus . That Tetracotylus c o ryphi c ephalus does not belong 
to the same genus with Proteocephalus f ilicollis has been shown 
by me in the descriptive study of the former species (pp ) 
of this article. The availability of the name Tetracotylus has 
questioned because of the name Tetracotyle Filippi (1854). These 
two names are not spelled the same and are therefore not homonyms. 
Braun (1894-1900) suggests that Tetratcotylus has been used in 
in place of Tetracotyle . If such improper use has been made of trie 
former name prior to the time when I.:onticelli proposed it then 
Tetracotylus has been rendered unavailable. I have not found 



evidence of this use hence I shall retain the name Tetrocetylus 
Mont io ell i (1891) with T. coryphicepl-ala Llonticelli as the type 
species . 

Habitat ; In Siluridae. 

The characters of the genus Tetracot ylus have been stated 
(p. ) of this work. 

The position of this genus in the order Tetraphyllidea is 
is difficult to determine. It apparently does not belong to any of 
the families as new defined therefore I suggest for it the family 
name Tetroco tylidae t with the characters of the type genus 
Tetrocotylus , are synonyms and hence the earlier name should be 

The question remains whether the name ffiroteocephalus is 
available. That name has been objected to on account of the name 
Prot&ocephala suggested by de Blainville (1828) for a cestode 
family. These two names are not certainly homonyms and the use of 
a name to designate a family does not invalidate it for use as a 
generic name. 

I, therefore, retain the name Proteocephalu s Weinland with 
Taenia f ilicolLis Rud. as the type. 



Having determined (pp. ) that Tetracotylus cory - 

l icephalus Llonticelli cannot belong to the family Proteocep - 
halidae because of its peculiar organization. I shall re-define 
the family P r o t e o c e phal i d ae thus: Genitalia in general as in 
other Tetraphyltidea. One or more pref of»irv ed irregularly alter- 
nating uteYq:- openings. Genital pore marginal. Vitellaria lat- 
eral, follicular, follicles grouped about central conducting ves- 
sel. Ovary b«-lobed, posterior, Cocapt , ootype , shell gland, 
uterine passage present. Uterus with lateral outpocketings , Vit- 
ellaria, testes, ovary and uterus within inner longitudinal muscle 
layer. Seolex with four suckers. Fifth sucker functional, ves- 
tigial, or entirely lacking. 

Habitat: in fresh-water fish, amphibia, and aquatic 


In this family belong the genera: 
(l) Protooc cphalus V.einland (1853). 

Characters of family. Head globose or conical, flattened 
dorsoventfally. Ilo^stellum. No spines or hooks. Ho fold of 
tissue encircling base of head or enfolding suckers. Suckers circul 

or oval. Fifth sucker functional or vestigial. Testes in a 
broad field betv/een vitell oria . Parenchyma v/ith close meshes. 
Musculature well developed. Eggs with three membranes. 

Habitat: Fresh water fishes. 


Type species: Protcocepha lus fil icollis Rudolphi ~ 
Proteocephalus g mbip;uus Dujsjtdln. 

(2) Choanoscolex. Hov. Gen. 

Characters of family. Head conical with a fold of 
tissue at the base partially covering suckers. ITo irostellum; 
no hooks * no spines. G-enital organs as in Pr oteocepha l us 

Habitat: Siluridae 

Type species : Choanoscolex abscisus (Eiggenbac&) 

(5) Corallobothrium Fritsch (1886). 

Characters of family. Scolex with four suckers situated 
on the flat anterior face of the head. Many irregular folds 
and lappets about margin of anterior surface may enclose suckers 
as in a coralla. No rostellum. Ho hooks nor spines. ITeck broad, 
short . 

Habitat : Siluridae 

T ype Spec ies; Corallobothrium solidum Fritsch. 

(4) Crepidobothrium Konticelli (1899) 

Characters of family. Head large, swollen, inversely, 
cordate, posterior margin interrupted and re-entrant into sucker 
cavity. Genital apertuwe irregularly alternating. Vaginal opening 
dorsal to cirrus pouch. Vagina anterior or posterior to cirrus- 
pouch. Testes in two lateral fields anterior to ovary. 
Habitat : Boidae , South America 
Type species : C repidobothrium Gerrardiv (Baird) 


(5) Acantho taenia von Linstow 1903 

Characters of family. Scolex rather small. V.'ith four 
rounded suckers. Apex of head conical in which is situated a 
vestigial. Fifth sucker, no rostellum. Cuticula of head and 
anterior part of "body covered with minute spines or "bristles. 
Segmentation, indistinct. Ho overlapping of segments. Testes 
in two lateral fields anterior to ovary. Ovarian lobes may be 
branched. Vagina anterior or posterior to cirrus. 

Habita t : in Varan idae 

Type species : Acanthotaenia shipleyi von Linstow 

(6) Ophio taenia Uov. Gen. 

With characters of family. Head globase or somewhat 
or somewhat tetrahgular. Ho rostellum. Ho hooks or spines. 
Suckers circular or oval, with margins entire. Fifth sucker 
vestigial. Heck usually long. Testes in two longitudinal fields 
anterior to ovary. Vagina anterior or posterior to ciijjis-pouch. 
Ovary flattened, sometimes alate. Parenchyma with fiyie meshes. 
Musculature weak. 

Habitat : In aquatic snakes, Crotalidae and Colubridae. 

Type species : Ophio taenio perspicua Uov. Spec. 

In the descriptive section the greater number of species 
of the above named genera have been described, with exception 
of species of Ac an t h o t ae n i a and GoroLloboth r ium. 

It has been impossible to write up the descriptions of 
certain species inquirenda of which no material could be ob- 
tained, and for which/only very meager descriptions have ever beon 


written. It is hoped that these may "be properly considered at 
another time. 


The cestodes of this family are constructed upon a 
uniform plan, which has been discussed very thoroly "by Monticelli 
(1891), Xraemer (1892) Riggenbac* (1896), Benedict (1900), la 
Rue (1909). This plan has also been reviewed in the descriptions 
of several species in the body of this work. It seems necessary 
to call attention here only to those structures upftn which now 
light has been shed by recent investigations. 

The external genitalia are situated on the margin of 
the proglottid as in other members of the order Tetraphyllidea . 
The inner genital organs are always situated within the inner 
longitudinal muscle layer. Only the cirrus-pouch, the vagina, 
and the ventral outpocketings of the uterus which become uterine 
pores extend outside of this muscle layer. Such a condition 
differs greatly from that found in Tetracotylus in which the 
testes, uterus and the vitellaria (fil.'Y Figs.tffr?/ ) lie wholly 
outside of the inner longitudinal muscles while the ovary is 
partially outside of that layer. A study of the literature does 
not furnish a parallel to^ relations of these organs as fomd in 
Tetracotylus . 

A fifth sucker is Ibund in a number of species of 
P r o t e o c e phalu s . Here it plainly has radial muscle fibers and 


appears to be capable of functioning as an organ of adhesion. 
Hot all Proteocephalids possess such a sucker. In such species 
a vestigial structure may usually bo found. 

In the tissue of the head of 0. f ilaroi dea just below 
the apex (PI. » figs.//f^} this vestigial organ occurs 
which in my former article. (La Rue . 1909; 24-26) I called an 
end-organ'. In that place I called attention to the fact that 
this organ was not a thickening of the tissues at that point. I 
also stated that no opening could be observed. These observations 
were made on the adult head. 

I further strted (La Rue 1909; 25) that the structure 
could be seen with great clearnesL in the head of the pi ero cere oils:: 
of this species and that in an unknown plerocercus from Ml - 
c ronterus dolomieu . Lac. it was a strongly developed.. I 
also pointed out the fact that in the Proteocephalids in which 
this organ was found there was no fifth sucker and that in those 
species in which the fifth sucker was found no vestigial organ 
had been seen by me or reported by others. 

In regard to the occurrence of this structure in other 
cestodes I quote the words of the former article (La F.uc ,1909 ; 2G) 
"Furthermore, it would be of interest to find out whether we have 
here to do with a cestode structure common to many unamed 
species in different genera or whether it be a structure found only 
in ITorth America. Apparently it has never been described in 
snecies occurring outside of Uorth America. " 

oince writing the above I have ezamined a large number 
of Proteoc .phalid species. In the genus P r o t e o c e phalu s 


(S. str. ) I have found it only in those species which have 
no fifth sucker. These are P. amblopliti s and P. macrocephalu s. 
I have thus far failed to find it in P. to yt d-oso. ^ P . per plexus . 
and ?. sin^ulari s . Among Ophiotaenia I have found it in 0. 
f ilaroides . Lofcnberg& and , 0. perspicua . I have been unable 
to section heads of any other species of the genus. This structure 
has been reported by Johnston, T.H. (1004: 104) in A c an t h o t a e n i a 
tidsv/elli , Johnston as : 'an spical muscle plug, an organ rarely 
met with in cestodes." So far as I an able to ascertain this 
structure does not occur in any species of Corollobothrium . I 
have examined preparations of a species of this genus in Dr. H.B. 
w'otrds' collection but have been unable to find this structure. 

In all these species in which this structure has been 
found there is never a fifth sucker. There is moreover no 
structure which one could rightfully call a rostellum. A true 
rostellum to the best of my knowledge does not occur in the 
Pr oteocephalidae . 

An examination of Brflnin (1894-1900) in regard to this 
structure shows that it has been found in a considerable number of 
widely separated species. Moreover it has never been reported 
in cestodes with a functional fifth sucker or a true rostellum. 

A careful examination of some sections thru the head of 
plerocercoids of this genus reveals the structure of the end-organ 
or muscle-plug. In those case:: this organ is seen to be an 
hypertrophied fifth sucker in which the tissues are packed with 
granules. The radial muscles have not yet disappeared but they 
are somewhat disarranged. The basement membrane is yet present, 
so also are the circular fibers whi ch show as cut ends. The 


openings to the exterior still remains. These conditions are 
delineated from "both transverse and frontal sections of the h 
head. (PI. b> flgBMM,)l(>J/7) 

A condition of the fifth sucker quite similar to this is 
found in the heads of Crep i dobothrium Gerrardft '. 

Here the sucker (PI. Figs. ) tho greatly re- 

duced in size and' tho it has lost its radial musculature re- 
tains its basement membrane and its opening to the exterior. The 
latter is only about 0.030 r.gto. in diameter and but a little 
deeper than that. 

The fifth sucker in this species evidently never 
degenerates as far as it does in some other species. It forms 
a transitional stage between the functional fifth sucker on the on<i 
hand and the extremely vestigial apical sucker on the other. 

Hot sufficient data have been secured to determine 
whether the fifth sucker is a rudfirontary rostellum. 


External measurements ■ Benedict (1900; 539) calls 
attention to the small dependence that can be placed upon ex- 
ternal measurements . It is certainly true^such measurements 
alone may be very misleading. Yet external measurements may prove 
of considerable value in/such variable forms as cestodes if they 
are properly taken. Such measurements when accompanied by a good 
description of external appearance form a very essential part of 
the species description, and should never be omitted except un- 
der the most unusual conditions. 

In making this study I have found that measurements of 


the head and of the suckers, together with the measurements of 
the strobila may sometimes furnish clews to the exact identifi- 
cation of the specimen. Measurements of the proglottids in the 
various regions of the strobile^ are of great assistance. 


Outline drawings of proglottids, "both mature and ripe, 
are of utmost importance. Such drawings, which show the relations 
of the male and female organs should aceompany the specific des- 
cription. Statements concerning the number and size of testes 
and variations in the same, number of uterine pouches, character 
of vitellaria and ovary, relative positions of vagina and cirrus, 
character and length of cirrus-pouch, and character of structi:res 
within the cirrus-pouch are of considerable value diagnostic 
value. Presence of muscular elements and cilia are aleo of 
value . 



In this section I have considered the species of 
Proteocephalus . Choanoscolex ilov. Gen, Ophiotoeni a IIov. G-en.^^</ 
Tetracotylus in the order named. It has also "been necessary 
to point out the true systematic position of other species which 
have heen mistakenly placed in this group of cestodes. 


(Pi. 3 



0. F. Muiier 

, 1782 


170b : 





17 1 ju : 

Taenia gasxerostoi 





Alyseimintnus gasterostei 

Goeze , 




Taenia liiieollis 

Poidolphi , 

18U1 : 



Halysis gasterostei 

Zeder , 




Taenia liiieollis - 





Taenia i'ilicollis 


laiy : 



Taenia i'ilicollis 





Taenia liiieollis 

Du Jar din, 




Taenia amoigua 

Du jar din, 




Taenia riiicoiiis 




luou : 

Taenia ambigua 





Proteocepnalus ambigua 

e inland, 




Taenia ambigua 




18tiv : 

Taenia liiieollis 


I8uv : 



Taenia iiiicou is 

Gui Scnneideriyu>; : 

84,86, 87. 


Icntnyotaenia ambigua 

Gui Scnneiderlvuo: 


iyoy : 

I cnthyo taenia ambigua 

luhe , 

1909 ; 



SPECIFIC DIAGNOSIS.- Characters or the genus. Very 
snail cectodes. Length up to 3b mm. Dreadtn up to O.b mm. 
Progiottids lev/, ib-2b. Segmentation Indistinct. Progiottids 
attached by tneir full width. First progiottids broader than long, 
u.^u mm. broad by u.uvb-u.uuu mm. ±ong. Mature progiottids about 
u.lti mm. long by U.37U mm. broad, ripe progiottids U.48-0.b6u mm. 
long by u.4b mm. broad. Last proglottid tapered posteriorly O.y/b 
mm. long by U.4^b mm. broad. 

Head small u.uyu-U.l20 mm. broad or perhaps a little more, 
not always v:eli set off from neck. Suckers O.U4^-u.Obb mm. in 
diameter. Ho fifth sucker present. Neck to iirst traces of 
segmentation about l.u mm. long or according to Schneider some- 
times equal to l/4 length ol' the body. 

GENITAL ORGANS.- As in genu:: . Seaital sinus marginal 
irregularly alternating, situated near middle or segment. The 
sinus situated on a papilla rrom prominence. 

Male organs .- Cirrus-pcuch about U.13U mn. long by 
U.u3£. U.U4U mm. broad, reacning about i/3 -i/4 across the breadth 
or the proglottid. Cirrus-canal much coiled in cirrus-pouch. Vas 
deferens excentrtc. T stes in t..o layers, about 7b-yu in all. 

Female organs.- Vaginal opening situated anterior to 
cirrus-pouch. Sphincter vaginae weakly developed. Receptacuium 
seminis present. Vitellaria quite compact, follicular, in lateral 
rieids. Ovary bilobed connected by an arched, mid-piece. Lobes 
spneroidai, thiefc. 


Uterus in ripe proglottids with b-7-ti lateral outpoclretmgs on 
either side. 

EGGS.- Outer membrane U.UbU-u.uvb mm. embryonal cover- 
ing U.U32 mm., embryo itseli mm. 


Host. ! 

, iiOoaiiiy 

! collector i 


Gasterostetrs acuieatus 

Goeze : 

Goeze 1 

tt tt 


Rudoipni. : 

Rudoipni 2 

tt n 

: Berlin 


Diesing 2 

II Tt 

Paris : 

Dujardin : 

Du jar din 2 

Tf Tl 


Beliingnam : 

Bellingham 2 

II 11 


Grimm 3 

Gasterosteus pungitius 


Du jar din 

Dujardin 3 

Tt rt « 

Halle : 


Diesing 2 

» t! 



Diesing 2 

rt ri 

: Keval Bst land 

;G. Schneider. 

G. Schneider 3 

It tt \ 

Pockhaia « 

levander : 

G. Schneider3 

; Finland. 

n tt , 

Upsala : 

Loiinberg : 

lonnberg 2 

1 J Reported under the name / ^lyselmir t nus gasteroste i. 

2) Reported under the name Taenia iiiicollis 

3) Reported under the name Taenia amblgua . 


HISTORICAL SUMMARY.- Rudolphi (181)1:114) gave the name 
Taenia ± ilioollis to the species or cesxode found by himself in the 
intestine oi' Gasterosteus oculeatus 4t Greifswald. Then he (1810: 
10b) gave a diagnosis and description with synonymy or this species. 
This synonymy snows very clearly that he himself considered his 
species to be identical with the forms described by eariier workers 
under the name of Taenia gasteroste iand Alyyelmintiius gasteros tei , 
Rudolphi' s diagnosis and description of his taenia fiiicollis 
are quoted: 

"Taenia fiiicollis R.- Taenia: capite subgloboso discreto . 
coiio longissirao . f iiir ormi . articuiis . ovariisque quadrat is , 

Muller im Naturrorscner St, IB, p. 28, Tab. 3. rig. fa, V. 
Taenia ex intest. Gasterostei. 

Batscn 13andw. p. 241. n. 14. 

G777€/ . Syst. Nat. p. 30'/y. n. 81. Taenia gasterostei. 

Goeze apud Zederum in huj. Ilachtrag p. 2t>5» Tab. 3. fig. 

1-4 Alyselmintnus Gcsterostei, 

Kudoipni in '.vied. Arch. III.l. p. 114. Taenia fiiicollis. 

Zeder llaturg.p. 354. n. 10. Haiysis Gasterostei. 

Hab. In Gasteroste i acuieat i intestinis Goezius Augusto, 

: ipse aliquoties sed capite orbam, Iunio, reperimus. 

Descr. Vermes duos tresve ooliices longi, fere lineam 

lati, candidissimi. 

Caput subglobosum, discretum, osculis orbicularibus , 

majuscuiis, binis tarn superioribus , quam inferioribus. collum 



filiforme, longissumum. uor^pus planum articuiis anticis minoribus, 
reiiquis subaequalibus, quadrans, ui timo roiundatQ- Ovaria quad- 
r mgularia, anguiis acutis produeiis, opaca, ut articulorum t an- 
il/ 1 partes inter anguios sitae peiiucidae sint. Vermis inde as- 
pectus iepidus, inter <juosiibet enim articuios maculae peiiucidae, 
et simul canalis lateralis peiiuoidi species oboritur* Foramina 
non visa. 

Obs.l. Goezius, qui ovaria non vidit, vermem proprie non 
ariicuiatum esse in posthumis reliquit, ipse vero hanc speciera aeque 
ac uiiam aliam articuiai5?n observavi. A Bothriocepnalo solicio, in 
Gasterostei abdomine hos r itante, omnibus prorsus noiis cliversissima. 
0bs.2-Yermem in aqua fitsviatili biduura vixisse, in puieali enecari. 
Goezius observavii. 

In his "Synopsis" Rudolph! ( 1819:148) quoted his diagnosis 
yf 1810, and habitat of the worm he wrote," In intestinis Gastoroste j 
aculeati Goeze August o, ipse Junis (Gynhioe) invenimus". 

Bellingham (1844) in his catalogue of Irish Entozoa of 
Taenia f ilicollis in the intestine of gasterecte us aoul e atus. His 
observations added nothing to our knowledge of the structure .of 
the worm. 

Dujardin (1845:583) reported that he had found Taenia 
f ilicollis in Gasterosteus aculeatus at Paris in 1838, His dis- 
cription, a part of which seems to be derived from Rudolphi (1810) 
reads: " — Long de 50 a 80 mm, large de 2 mm environ; — tete pre s que 
globuleus, distincte et portee par un cou tres-long, filiforme; — 


sans trompe; — articles presque carres, contenant dcs ovares (?) opa- 
ques egalement carres, qui laissent entre eux, et sur les bords, des 
intervalles deni-transparents.** 

Rudolphi et Goese l'ont trouve en Allemagnc, dans le Gas- 
terosteus aouleatus . Je l'ai trouve aussi dans ce poiscon, a Paris, 
en 1838; j'y ai vu des aeufs a double enveloppe, dont l'externe, 
mucilagineuse, est longue de mm, 06 a mm, 10; l'enveloppe intern 
globuleuse, est large de mm, 036, et les crochets de l'embryon 
sont longs de mm, 012 

Dujardin (1845:583) also described a form from Gasterosteus 
pungitius (leavis) under the name Taenia anbigua . His specimens 
were collected at Rennes, France • His description reads: w — Long 
de 6 mm, large de mm, 5 a mm, 8, forme de uuinze a dix-sept 
articles peu distincts et de forme tres-variable ; — tete petite, largi 
de mm, 17, sans lrompe et sans crochets, tantot retractee, tantot 
saillante, quelquefois globuleuse et separee par un cou tres-etroit, 
quelquefois a quatre lobes distincts, correspondant aux ventouses 
larges de mm, 068 a mm, 07; — cou tres-contractile et dilatable, 
traverse par quatre canaux larges de mm, 009; — premiers articles 
malec avec les orifices genitaux irregulierenent alternes, et les pe- 
nie longs de mm, 16, larges de mm, 032, rides transversalement ;- 
derniors articles informes, remplis d'oeufs globuleux a douole en- 
veloppe; — enveloppe externe mucilagineuse, longue de mm, 053 a 
mm, 026, avec des crochets de mm, 0095. n 

Je l'ai trouve plusieurs fois a Rennes dans l'intestin du 
Gasterosteus laevis . Son nom specifique exprime sa ressemblance ave< 
les scolex et les caryophylles*" 


In Dujardin' s description of his T. f ilicollis and his 
!T. ambigua he compared, it will he noted that in the measurements 
of the second egg* membrane there is very good agreement • The 
variation in the siae of the outer egg membranes in Proteocephalus 
is large hence but little value can be given that measurement. 
Unfortunately Dujardin did not give the measurements of the embryo 
in either case so no comparison is possible in that respect. 
Dujardin records a difference of 0.003 mm. in the length of the 
hooks of the embryo f That measurement however is difficult to 
make with accuracy and much weight should not be given this 
character as a merns of differentiating the species. 

It is probable that Dujardin' s specimens from Gasterosteus 
oculeatus and from G. pungitius b elonged to the same species. 

We inland (1858:53) made Taenia ambigua Dujardin the type 
of his genus Proteocephalus and with that species he included 
Taenia f ilicollis and Taenia dispar . 

Diesing f 1850: 512) recording Taenia filicollis F.ud. 
added nothing to Hudolphi's diagnosis and description of that spe- 
cies. He gave a complete synonymy up to his time. He stated that 
this species had been found in the intestine of Gasterosteus 
aculeatus in August by Goeze and in June at Greifswald by Rudolphi; 


in sepxember at- Berlin by de Oilers; at Paris by Du jar din; in 
Gasterosteus pungixius ax Kalle by ITixsch* ax Greil'swald by Creplin. 

On Taenia ambjlgua Dujardm, Diesing (18bOJ gave nothing 


Grimm ( ld'/2: 243-^46 J i'ound a iorm in G asxeros^eus 
aculeatus which he identified as T aenia afabigua Dujar^in. Grimm 
published no drawings oi his specimens. The import***" parts 01 
his description are ijuoted here. 

TAENIA AMBIGUA Duj. — Im Darme des bei uns so haui'ig 
vorkonrnenden Stichlings (Gasterosteus aculeatus ), von denen ich 
im vergagenon Sommer mehr als 100 aul'geschnitxen habe, fand ich 
unter andorn 6 Exemplare oiner Bandwurmga''; ' . wnfttao*; older, 

sich sehr ieichx von alien ihren Yorwandten und sind augensci ein- 
lich zu der von Dujardin aui'gestellten Dpecies Taenia ambigua zu 
rechnen, obleich die Bestimmung, einer Kurzen Beschreibung des 
auaseren Habitue nach, hochex schwierig isx und sehr ieicht zu einem 
Irrfhorn. fi).hre-n. K^nn. 

Das gro'sste von mir aufgei'udene Exemplar hatte 30 Mm. 

Lange und IMm. Breixe. Die Gossenverhaltn:2sse der einzeinen 

^angen exemplars sind focgene,: die lange -iLer Xoopchens 0,' Mm 

Korphertheile eines ii t 5 i.Im. dessen Brer -3 — 0.25 mm., dessen 

Dicke u,it> Mm. , der Durchmesser der Saugnapfie — 0.0b Mm* der 

Lc!nge des Haises — 1,1 Mm. , L.'L^- aor ersxen unr'eiien Sliec 

t ui4 Km.; dieselben verlangern sich allmaiig, indem sie u.02 Mm., 

u.03 Mm. u. s. W. lang werden; reil'ere Glieder. ungei'ahr aus der 

MiXte des 77urrnB t haben eine Lange von u.20 Wty und das Vollkoramen 


reil'e vorletzte Glied ist 0,43 Mm. long. Die Dicke des Wurmkorpers 
steignt bis skl 0,5 Mm. 

Das Koi.'i'ciien des "tfurms von der Seixe betrachtet, erschoint angesch.vollen; wenn man es von der Flacfre betracltet, so sielrc 
man an den in Weingeist conservirten Exempiaren oine Furcne, die uber 
den Kppi, an dessen i'reiem Ende, von der enien 1'iacnen Seite des 
V/uraa zur andern verlauft. Diese Furcne ist aber gewiss keine con- 
state Bildung, wird aber dadurch hervorgeruien, dass der Wurm^ 
uDerhaupt je ir contract il, sein Kppi'chen biters einzieht, namentiich 
wenn er in Weingeisx geiegt wird. iiiner Kiissei besitzt er nicht. 
Die Saugnapie, die eine runde Form haben, sind paarweise auf den 
ilacnen Seiten des Korpers entspreclienden Xopftiieilen angeordnet. 
Die Geschlecnxsbf fnungen iiegen am Rande der Glieder, eine uber 
der andern. Alie Glieder, die mix dem Alter allra'ahfig breiter werden, 
haben eine ziemlich unregelmassige Form, besonders aber inre Rander. 
Die Far be des Wurras isx rein v/eiss. Indem wir noch hinzusetzen, dass 
das te ste Glied an seinem hintern Rande bestandig et'.as aui'gescJi- 
iixzt is-c, naben wir eine raagiichst vollstandige Darsteliung 
des Aeussern des Bandwurms geliei'ert. 

Dieser Bandwurra isx .o^nst zart - 

! f 

Von dem we ibli often Go sciilsciixs organ nabe ich nur die 
Uterus aufgefunden : die 
Keiradru2( Ke^jnirVi -legt in der Mitt e des Gliedes und hatte in dera 

unttersuci "cen stadium ein spindeilormiges Aussehen; von inr geftz 
der Jterus aus, der bus einem Rohre mix iemen raserigon 
Wandungen oesxent, das mehrere Schlingen Bildex. Der uxcrus veriaiiit 


in die Scl^ jide, die sich zura Seitenrande des Gliedes begioDt, 

wo sie nacn aussen mit einor ziemiicn gVossen oefi'ning miindot. 

Ileben aem weibiichen Geschlechtsorgan liegt eine andere 

oeffming, die in das mannliche Gesciilecntr per**? xuhrt. Das lotzte 

besteht au^dem Koibenformigen Cirrusbeutei mil; dem nach innen 

gewendeten Cirrus, der an seinem .Ende etwas anschv/illt. Hinter dem 


Cirrusbeutei liegt die doppelte Samendruse. 

In dera letzeten, also am meisten entwickeClten Gliede waren 
nur Keirae, nicht aber entwickelte Sir zu sehen. 

Von Linstow (1878: £31 J reported as hosts for Taenia 
iiiicollis and Taenia a mbigua only those hosts that had been reported 
by Diesing (l8bo) 

Zschokke (H J4:lb-17J reported and briefly descrioed a 
form from the Percr f luvitilis Lake Lucerne which he corsidered to 
De Taenia f ilicollis Rud. His forms has "been identify as belonging 
to P. dubuis N.sp. 

Leidy (1886) though that he had found Taenia f ilicollis 
in the intestine of Annia caiva from llorth Coralina. This was a 

mis determination. His species was probabijr the same as the species 
for which I have proposed the name, P roteoceohalus perplexus II. SP. 

Lqpnberg (i88y:lb) reported e finding of Taenia 
iiiicollis in Gasterosteus p ungit- ' (/$ it Ups^Ia. He added very 
little to Rudoipni 1 s- diagnosis. 


Kraerner (l8y2) reported and described a form from the 
intestine 01 Coregonus f era from Lake Lucerne, which he i'irst 
identified as Taenia f iiicoilis Rud. In the same paper Kraemer 
showed that his Taenia i'ilicollis and his Taenia oceliata were 
identical. Riggenbach (18y6) accepted Kraemer 's determination. 

This species has been shown to differ from the taenia 
1 iiicoilis of Rud. and is described under the name Proteocephaius 

■ii I, i I, iii i *- _ 

f allax n.sp. where a full discission of the position of Kraemer' s 

species may be found (p. ). Benedict (lyOOJ described as 
American species of cestbdes under the i xme Proteocephaius 
f il icollis (Rud. ) In a note he changed the name to P roteocephalus 
ocellatus (Rud.). }Je accepted Kraemer' s statement that Rudolphi's 
Taenia oceiata a nd T.f iiicoilis were identical. For this form 
which proved to be a new species I have proposed the name, 
Proteocepnalus exiguus . N. sp. 

Schneider (1902:21-^3) reported a form from Perca 
fluyjctilis . in Finland, as lohthyotrenia iilicollis Rud. This form 
he later (iyu3:l3 and iuub:lb-l7) considered to be identical with 
0. F. Miiiler's Taenia Percae .. Schneider ( 19u2:84,8b&88) reported 
a specimen from Gasterosteus pungitiu s as Iohthyotaenia f iiicoilis 
I Rud. In a later paper (lyub: 21-24) he consider this form to be 
! iohthyotaenia ambifrua Du jar din and here he gave the first good 
description of this species. 

Luhe (lyoy) gave a diagnosis of this species under the 
name Ichtiiyotaenia a m b igua Dujardin. His diagnosis added nothing 
to the data given by Dujardin (1845) and Schneider (lyUbJ. 


DISCUSSION.- There is no doubt but that Grimm's and 
Scnneider's specimens belong in tne same species with Taenia 
ambigua Dujardin. However, a question arises whether Dujardin' s 
Taenia ambigua is identical with the earlier Taenia gastcrostei 

( T. rilicollis Rud. ) . It is no proor of the identity of the two 

one. host 

species that they are round in an idividual host or in .ecies, 
numerous instances might be cited in wncih a species or an individual 
in the species served as host ior two, or more, closely related 
species or cestodes. Nevertheless knowledge of the host is recognized 

of- gre<o-* 

to beAimportance in proving an identity and rrequently the knowledge 
that two rorms are from the same host species i'urnishes the first 
intimation that they may prove to be identical. Again the lac: 
tnat two rorms similar in outward appearance have been taken rrom 
unrelated species nay give an early intimation that the rorms belong 
to different species. 

The fact that some parasites seem to be restricted in 
habitat to a single host species might be ad/anced here as an 
argument against the identity or Taenia gasterostei and T. ambigua . 
In reply it may be said that in general closely allied hosts having 
a 7/ide distribution and inhabiting tne same regions and especially 
wnen they nave similar leeding habits very rrequently harbor the 
same species of parasites. Gasterosteus acuieatus and G. pungitius 
come under this description. T. ambigua h.. s been reported rrom 
eacn ot tnese species so also has T_, gas oerostei (i^iiicoiiis •) 
Grimm's specimens or T. ambigua came i'rom G. aculeatus while the 
specimens or Schneider and o± Dujardin were rrom n . j>unnTius f 


lion neve been seated before to belong in the same species. The 
specimens of T. f ilicoliis which Rudoipni, he Oilers, .Dujardin 
and Bellingnam reported were round, in G« oculeatus wniie the T. 
iilicoiiis reported by ITitscn, Creplin and Lo'nnberg were round in 
G. pungitius . Tnis state 01 arrairs causes me to believe that T. 
ga s teroste i ( iilicoiiis ) and T. ambigua are identical. 

The most serious argument against this identity is tne 
fact that Rudoipni's description of f iliooilis idicates a 
larger form than does Dujardin' s description of T. ambigua . It is 
however a v;eii known fact tnat a considerable variation in length 
and breadtn may exist witnin the same species of cesi;odes. These 
variation may be due to tne degree of maturity and. to the states of 
contraction of tne idividuai worms, it is not kno" n tnat Dujardin' 
' specimens of T. ambi gua nad attained tneir full development mor is 
I tnis known concerning the specimens of Grimm or Schneider. 

Tnere is not suificient evidence at haftdto prove a 
i positive identity for T. gasteroste i and ambigua . The evidence 
, for tne identity nowever is stronger than that against it. There- 
; fore in tnis work I shaii consider T. ambigua Dujardin to be 
identical witn T. gastefcotci (Iilicoiiis) and that the name T. 
ambigua is a sjnonym of the earlier name T. gasterostei . The latter 
specific name then should be retained and in the lignt of our 
present knowledge would be known as Proteocepnaius gasteroste i( 
MATERIAL.- G. Schneider's material was found for the 
most part in ^astoros teus pungitius tne Qlbersee near Reval 


costodeu ojf 

in Kstland in the months oi ^ay and June, liever more than lour 
specie? were found in a single host. He aieo examined a single 

specimo . round "by Dr. K. M. Levander at ?or£haia, i'iriand in 


Gasterosteusfcungitius reported by Schneider ( ±yu2:B4,«b*yy) as 


Iohthyotaenia f ilicoilis I Rud. } 

For my study Dr. H. B. 77ard very kindly secured a specimen 

oi" this form stained in carmine and mounted in toto, from Dr. K. 

M. Levander in Heisingrors. Tnis was one oi Schneider's specimens, 

It bears the laoei , "ichthyotaenia ambigua int. Gast pungitius 23/1U 

VI U4. 

The description which follows is cased on the description 
of Schneider ( lyu5:21-h;4 ) and on my observations on this one 
mounted specimen. 

EXTERNAL CHAKAC'JfEKS • - Schneider's longest specimens 
measured up to 3b mm. long by a max. breadtn or O.y mm.. The longest 
specimen had 17 progiottids. (If these the anterior progiottids were 
broader tnan long. Those are joined together by nearly the whole 
breadth. The progiottids that are filled with eggs are longer than 
broad and have a rounded sac-lilce form. 

The specimen^ observed by me A about 2b progiottids. Of 

this number there are only a few young progiottids. The number of 

is considerably larger aniL— the numb r of proglottid: 
mature ana* nearly mature progiottids with eggs in the uterus is 

again small. The strobiiation is indistinct. The first progiottids 
measure about u.U7b-u.U«o mm. long by y.2uO mm. broad, the mature 
progiottids about u.i8U mm. long by U.37U mm. broad and the ripe 


above 0.48-0.660 mm. long by o.4t> mm. broad. The last proglottid 
which is tapered posteriorly is o.y'/b mm. long by o.4^b mm. broad. 

The scolex recording to Schneider is small, o.loo-o.lEO 
mm* broad, not sex ofi well from the neck. The suckers are 0.055 
mm. in diameter. No fifth sucker is present. The neck is relatively 
bread and its length is about one-fourth the "body length. 

Ite the specimen examined by me the breadth o±' the head 

at the base oi' the suckers is O.Oyo mm. The head, is very short and 


not set off well from the neck which is comparatively broad. The 
suckers are about 0.042 mm. in diameter over ail. The neck back to 
the first traces of segmentation is about 1.0 mm long, about i/8 
the length 01 the entire body. 

Excretory s yst em.- Schneider found that the excretory 
vessels are quite wide; that the two ventral vessels which communicate 
in eacn segment with the exterior by a subraatfginai pore are U.U3 mm. 
in dimeter; that the dorsal vessel are smaller with a deameter of 
U.UU75 mm. 

GENITAL ORGANS.- The genital sinus is marginal, 
irregularly alternating, and situated about the middle of the 
proglottid length. The genital sinus is rather prominently situated 
on a little eminence which is most evident in ripe proglottid^. 
It may be due perhaps to the pushing out of tnat region by near- 
by organs. , , 

MALE ORGANS. - The cirrus-pouch /v is about 1/3-1/4 as long 
as the proglottid is broad. Its length is about 0.130 mm. by 0.052- 
O.O40 mm. broad. 


The cirrus-canal according to Schneider is greatly colled within 
the cirrus-pouch and it broadens out m its proximus end to form a 
vesicula U.U3U ram. broad. The coils or the deferens outside the 
cirrus-sneatn are excentric i'or the most part on that side of the 
segment m which the genital aperture occurs. 

40-bu in a layer, Since they are apparently in two layers their 
total number is probably about 75-yu. Schneider round that the 
testes measure U.U55-U.U6L' mm. in sagittal and U.U35-u.U4t> mm. in 
freHtal diameter. / 7 i 

Female organs .- The v* rina^opens anterior to the cirrus- 
pouch. Schneider found a weak spAincter vaginae near the opening 
of the vagina. The vagina passes in a curve to the middle of the 
segment and thence posteriad to the interovarial space. He found 
a small receptaculum seminis just anterior to the ovary. 

The vitellaria are quite compact f oiliculaTmasses in the 
lateral fields of the segments. 

Of the ovaries Schneider writes 1 / The ovaries appear as 
spherical bodies when seen from the surface. In the posterior 
segments after they have filled tne uterus with eggs, they wither 
cor lately or the remains of the genital organs, pressed together 

//ito a triangular area, are recognizable as two small round bodies 

I hoid thai Grimm \ld'/z p. 246) erroneously considered this 
triangular area to be the ovary when he says: "the ovary lies in the 
middle of the segment and has in its observed condition a spindle — 

I found the testes in the toto preparation to be about 

in the posterior region of the segment." 


like appearance." 

The ovary is bilobed, the lobes collected by a short arched 
midpiece. The iobes are spn roidal. In the ripe proglottids observed 
by me the ovary could still oe distinguished. 

Schneider round that the uterine passage opens into the 
uterus posterior to the middle oi" the pooglottic opposite the place 
wnere the performed opening oi" the uterus is found. 

The uterus which is a median tube in young proglottids has 
b-7-b lateral outpocketings on either side in ripe proglottids. 
Schneiders drawings show 3-8 oi these lateral outpooketings on either 

2GG3- As observed by Schneid ir the outer lnembrare- oi* the 
egg measures u.uou-J.UYb mm. tno erab/yynal covering measa es u.u32 
jn. and the embrjyogi it sell U.U27 mm. 

In th i size oi' the head and in the size oi" the suckers 
Schneider's form is smaller than Dujardin's. In most other respects 
the two i'orms agree admirably. In the measurement of the embryo 
the agreement is very good. If Dujardin's measurement of the head 
were made on living specimens the difference between measurements 
ox Schneider and Dujardin could be readily accounted for. These 
differences are not of sufficient weight to preclude the possibility 
of the forms being identical. The very close agreement ox the 
embryo is or ror greater weight than the differences in the size 
of the nead and suckers. Schneider form and Dujardin's are to be 
considered identical. 


RELATIONSHIPS.- P. ambiguas somewha-c resembles somewhat 
P. adonis Barbieri but it may be differentiated rrom that species 
oy its relatively shorter cirrus-pouch, Dy different position of the 


mass of coils of vas dererens, A Dy its larger testes wnich are in two 
layers while the testes of P. agonls are in one layer. P. agon is 
has no genital papillr a while in P. ambiguus there is a simulation 
of a genital apille __P. am bi gnus is readily distinguished from 
P. dubuis ft, so ana P . fallax ?*.sp. Dy its smaller size, by its 
lack of a ^'ifth sucker, by its more numerous testes, shorter cirrus- 
pouch, and smaller proglottids. The embryos of P. ambiguus are 
smaller than those of ?. fallax * 

P. ambiguu sjdifTers from P. pusilius M&rd and P. exiguus 
h.sp. in not having a fifth sucker, in having smaller suckers, a 
smaller head, more testes^ and fewer uterine pouches. 

This species differs greatly from P. perca e in the lack 
of a fifth sucker, in size of body, head, and suckers, in the 
proportions of the proglottids f in the size or the cirrus poucn,ovarie 

This species is smaller than P. esopis , It has a shorter 
cirrus-pouch^ a larger number of testes. The proportions or the 
proglottids are difrerent. The two species are alike in not f o5 ~ 
s-essing a firth sucker. 


(PI. jt figs. (pi-7i) 

1884: Taenia yilloollis Zsebokke, 1884: 16-17. 

1884: Taenia ftcellata Zschokke, 1884: 13-14. in part? 

SPECIFIC DIAGNOSIS.- Characters of the genus. Small 
cestodes up to 40 mm. long by 0,80-1.20 mm. "broad. Segmentation not 
very distinct. Number of proglottids up to about 100. 

Head small, variable in shape, well set off from neck. 
Length of head 0.10-0.14 mm. breadth of same 0.127-0.212 mm. Four 
suckers 0.06^-0.080 mm. in diameter, cavities deep, sucker outlines 
round or irregular. Fifth sucker 0.026-0.037 mm. in diameter. 

IJeck slender 1.8-3.0-3.5 mm. long by 0.085-0.140 mm. 
broad. First proglottids broader than long, o.20 mm. broad x 0.050 
mm. long, i.iature proglottids quadrate broader than long, or longer 
than broad, 3.02 mm. long x 0.34 mm. broad to 0.595 mm. long x 
0.680 mm. broad. Ripe proglottids quadBate, broader than long, or 
usually longer-than broad , 1.19 mm. long x 0.680 mn. broad - 0.680 mm. 
lo£g 1.02 mm. broad. End proglottid, triangla^snpll , functional. 

GSITITAL ORGAIS.- Genital opening marginal, near middle 
of proglottid or anterior thereto, irregularly alternating. Uo. 
genital papilla. 

Male Organs.- Testes in two layers between vitellaria anter- 
ior to ovary 55-60 in number. Vas fleferens a mass of coils in mid- 
field. Cirrus-pouch long, slender e. 265-0. 370-0 .425 mm. in lengthx 
0.070-0.085 mm. in breadth. Ratio of length of cirrus-pouch to 


i proglottid breadth 5/11-1/2. Cirrus slender straight when protruded 
short and slender. Ductus e jaculatorius straight. 

Fein a le organs .- Vaginal opening anterior to cirrus-pouch. 
Sphincter vaginae weak, situs ' near opening of vagina. Vagina 
crossing cirrus-pouch near middle. 

Ovary Bilobed.- Lobes thick somewhat quadrate in nearly 
mature proglottid^ smoothly ovoidal in fully mature and just parsed 
mature proglottids. 

Vitellaria lateral, sparse, follicles small. 
Uterus with 7-10-12-14 lateral pouches on either side. 
Uterint pores one-two on ventral surface. 

-.GGS.- 7/ith three membranes, second membrane granular 
thick, 0.042-0.053 mm. in diamter. Embryo about 0.032 mm. in 

HABITAT : Intestine Perca luviatilis . Lake Geneva 
and per haps other lakes and rivers of Southern or Central Europe. 

HISTORICAL SUMMARY.- Zschokke (18 4:16-17) described some 
specimens of cestode found by him in Perca ^luvi ^tilis from the 
Lake Geneva. In so r :m respects his description agrees fairly well 
with P . ^allax n. sp. . from Qoregonus £era. However in regard to the 
ovary there is considerable difference. Moreevor it scarcely seems 
probable that Perca j flut idri iis and Coregonus ^era would normally 
harbor the same paras i rc speci es,Vlth this doubt in wind some 
material which Br. H. B. >Vard had reciived from Prof essor Parona anc. 
labelled Taeni a o oellata fro m Perca ^luviat ilis was carefully 


investigated. It was found that this material which at first 

glimpse appeared to he identi il with P» la/la x nss P» from 

Corerronu o fzm was after all sufficiently different to make it 
a new species. 

SOURCE Of "ATERIAL.- Alcoholic material in Professor 
Parona's collection labelled "Taenia Ocellata . Perca Fluviatilis 
" locality not knovn and slides and alcohplics of the same in Prof. 
H. E. Ward's collections. 

EXTERNAL CHARACTERS.- These cestodes are of small 
size, "being short and slender. The longest specimen observed by me 
measured about 4C mm. in length by about 0.80-1.2 mm. in max. 
breadth. In such a specimen there are about 30 proglottids with 
eggs in the uterus, 12-18 mature proglottids, about 3 which show 
more or less developed sex organs, and very few in which the anlagen 
of the sex organs cannot be seen, in all about 80-00 proglottids. 
The segmentation is not very distinct for the proglottids are joined 
by nearly their entire br adtli. The corners- are slightly rounded. 
Zschokke stated that the margins of the proglottids are not rounded 
and their angles are romc.h. t rounded; that the animal never presents 
a crenated appearance and that it has the ap earance of a ribbon. 

Zschokke found from 60-100 segments in the strobila. 
His longest specimen was about 4 cms. long and was about 1mm. bread. 

The head (Pl.-^ fig>» ') is small of variable shape. 
Frequently its anterior face is somewhat con: ?al, at other times 
flattened. The head is flattened dorsoventr ally. Its length varies 


from 0.100-0.140 mm. Its breadth from 0.127 in heads turned slightly 
sidewise to 0.Z12 mm. in slightly dictated heads, Ac its broadest 
part the head hears four sucker which are directed outward and up- 
ward. These suckers appear to he variously placed on the head 
depending on the state of contraction. A small fifth sucker 0,026- 
0.037 mm. in diameter is situated at the apex of the head. The four 
suckers vary in size from 0.069-0.080 ram. in diameter. They are 
usually nearly circular in outline but may be a little irregular. 
The cavity is deep andofirregular shape or it may be nearly round 
in outline. 

The neck is/slender, 1.8 mm. to 5.0-3.5 mm. in length by 
0.085-0.140 mm. in minimum breadth. The length of the neck was 
determined, by measuring from the hetd to the first visible segmenta- 
tion in stained preparations. 

Table showing cirnensj ons of neck, head and suckers in 

lie ok 

len gth 







length ' breadth 

0.12.-0.140 0.144 









0.069 : 0.069 
0.069; 0.069 

Fifth sucker. 



1) Heed slightly turned sideways. 

Zsehokke's description of the head, sucker and neck reads: 
"La "tote est petite, 

distinct e du cou, arrondie en-avant. Son pou^tcur est garni de quatre 
ventouses dont la position relative est fort variable ( V* PI- figs*) 

Ces organes de fixation Smt tre: pro^Dnds et puurvus 
d'loie puissante musculature. Quelauefois, ±{3 sont reunis suyune \ 
defaces, de m<|nciere quon peut lea foir de f ce to«s les quatre a 
la fois. Frettue»!»ent elles font saiUie au dela des pourtours dc 
la tete. Celle-ci prend alors une forme lobe'e on frangee fort variable 
; selon le degre' de contraction des ventouses. La tete presente ainsi 
i souvent des proeminences distribue'es regulier e3ment . Les ventouses 
peuvent in Sine affecter la forme de petites troupes irotractiles. 

Le cou est long, filiforme, ©'pod £ sfi' en-aririfje ; il n'occup 
cependant jamais plus dliw cini£uiewe ou ak>i quart de la longeur 
total? de 1 'animal, tandis que chez le taenia I ongicollis il 
at^einct ■ facilement un tiers " 

There is pretty good agreement between^description and 

mine . 

The first proglottids are A than long, 0.£0 mm. broad by 
0.050 m . long. These soon elongate somewhat and then the anla^gen 
of the genital organs begin to appear. The proglottids as they 
become mature elongate and also increase in width. They may be much 
elongated^ nearly square or broader than long depending upon the 
state of contraction. In the same strobila the following measure- 
r ' — — — . = 


mentsi#ere taken from anterior to posterior^ some fev; proglottids 
being omitted at intervals, the measurement ©/length "being stated 
first: 1.02 x 0.34 mm.; 0.68 x 0,485 mm; 0.595 x 0.425 mm; O.ST-Ox 
0.595 mm; 0.595 mm. x 0.68 mm. The last is the measurement of the 
l«st proglottid not having eggs in the uterus. 

Ripe proglottids may be nearly square, longer than 
"broad, or broader than long. They may measure up to 1.190 mm. long 
x 0.680 mm. broad , 0.680 mm. long x 1.02 mm. b:-oad, 0.680 mm. long 
x 0#850 mm. broad. 

The end-proglotti<3 is triangular in outline, the 
posterior end being rounded. It is 0.425 mm. broad by 0.476 mm. 
long. Apparently It "is functional. No excretory pore could be 
seen at its posterior end. 

Sschokke wrote of the proglot t ids "Lee premiers articles 
sont courts deux a trois fois plus larges cue lange # Les suivants 
deviennent carres et les derniers sont ordinairement plws long que 
larges. larticle terminal est awondi en arriere" 

GE1IITAL ORGANS.- Ihase are typical of the genus. The 
genital opening which .is situated near the middle or slightly anter- 
ior to the middle of the lateral margin of the proglottid alternates 
irregularly in the strobila from right to left. There is no genital 

Male Organs.- The test*s^are nearly round or at times 
slightly eompressed. They measure 0.063-0.079 mm. in diaieter. 
They lie. in two partial layers which cover the entire field 
between the vitellaria anterior to the ovary. Th€«V number is 


The vas deferens forms a small compact mass of coils 
in the mid-field of the segment. The cirrus-pouch ±9 long, quite 
slender, smooth in outline or at times somewhat constricted near 
the inner end by the contraction of circular muscle fibers. The 
cirrus-pouch measures 0.370-0.425 mm. 10ng in ripe pwylottids, 0.265- 
0.292 mm. long in proglottids with few uterine eggs bind .226-0,265 mm. 
in mature proglottids where the cirrus is being protruded. Its 
breadth is about 0.070-0.085 mm. It extends from 5,/ll to l/2 
across the proglottid breadth or in some cases it may extend slightly 
past the middle. 

The cirrus is straight and slender. When protruded it 
may extend about 0.1 mm. past the margin of the segment. The 

| protruded eirrus is/slender, or if but slightly protruded conical. 

j The ductus e jaculatorius is straight and of the same size as the 
cirrus of which it really forms a part. 

Female Organs. - The vagina A opens Into the common genital apparently anterior to the cirrusnpouoh. Hear its opening 
it possesses a small sphincter vaginae./ The course of the vagina 
is slightly anteriad then postoriad and -mas^d in a long arc which 
crosres the cirrus-pouch near the middle of its dorsal^side. Its 
entire course into the intepovarial space has not been traced. 
A receptacum sominis has not been observed 

The ovary is posterior and is biloberl. These lobes in 
nearly mature proglottids may not show the mitipieOce. The lobes 
may appear aS more or less irregular quadrate bodies in the posterior 


corners of the segment. In such a condition they much resemble 
the condition v/hich Zj3hokke delineated (PI* fig. ) , In 


more mature and ripe proglottids the lobes^seen tObe more even in 
contour (PI. tig. ), In ripe proglottids the ovary A have a span 

of 0.425 mm. The lobes are heavier and thicker than in 2.{a-£fajc 

- n*.3p. 

The organs of the interovarieJL space have not b^en observed. 
The vitellaria are lateral follicular masses. The 
follicles are small and not very compact. 

{jp/;if>-f-7°J \ 

The uterus A in ripe ' proglottids is made up 0/ 7-8-10-12 Or 
even 14 lateral pouches on either side, the more common number 
being 8 or 9. These pouches are more or less irregular in 

shape . From one to two ventjal uterine pores have been observed. One 
large pore oceujs quite regularly near the middle of the proglottid: » 
A smaller pore may sometimes be found farther anteriad. 

EGGS.- The uterine eggs have three membranes^, an outer 
AjCaline.of variable size v/hich was not measured; iL second shell 
thick' 'ranu(a\i/r and nearly spherical measures 0.042 and 0.048 — 
0.053 mm. usually about 0.048 minj And qjb. inner/thin membrane about the 
embryo. The embryo me asure$ about 0.03T: mm. in diameter. Vhen it is 
elongated it may measure u.026 x^ . )jD37 mm. or up to 0.032 x 0.037 mm. 
The average measurement is about 0.032 mm. 

Zschokke^ description of the genital organs reads thus: 
"Les orifices genitaux se trcuvent au fond d'un bourrelet circu- 
In ire, situee au milieu ou un peu audessus des bords l^teraux. 


L'alternrnce de la position a gauche ou a droite est lrregulier 
Les cirrhes sont courts, coniques, leurs poches etroitefl'i 
RUDOLPHI parle de ovaria qusdrangularia ; en effet le parenchyme 
est rempli de vesicules, qui dans les articles jeunes, ont, une 

forme ronde ou ovalaire et deviennent, dans les proglottis plus 

a r iff 

murs, carrees et anguleuses. En realite ces vesicules sont a 

/ ' i 

l'etat normal pyriformes; leur aspect carre est le resultat de 


leur pression ceciproque. Du rest ce ne sont pas des ovaires, 
ce sont des testicules dont chacun presente un petit canal 
defe'rent. Tous ces canicules se reunissent dans un canal ex- 
creteur commun. 

Deja DUJARDI5T, en citant la description de RUDOLPHI, Met, 
derriere "ovaires opaques, egalement carres," un point d' inter- 
rogation et DI.ESING n'en parle plus. 

Le vagin est court, il aboutit a une pcche seminale assez 
spacieuse. L ! uterus sous forme d'un tube large suit la ligne 
me'diane di proglottis et se rend en de'crivant quelques fabiles 
lacets vers le bord pcsterieur de l'article. La il decrit un 
noBud de lacets et se divise enfin en deux branches, qui, de 
leur cote, constituent a droite et a gauche une masse d'ovaires 
lobe's ou contournes. Entre les ovaires, tcuchant le bord pos- 
terieur du proglottis, se trouve la glande vitellogene. Des 


amas gl^ndulaires, peut-etre la continuation directe des vit- 
tellogenes, se trouvent le long des bcrds lateravx. 



RELATIONSHIPS . - This specios is much smaller than P. 
ercae (Mailer), It is most closely related to P. ^al^ax n^6p. 

from Coregonus ^ sra It differs from that species in having regular^ 
a larger number of test e s (ahout double the number to be f ound ( /P . 
^allax ) which are also larger, In__P. ^allax the testfs arc in 
one la£er ; in this species in tvrp partial layers. 

The cirrus-pouch in this species is usually a little 
longer than in P^ f*-IU y and it is relatively 

longer in proportion to the proglottid breadth. Therq may be more 
numerous uterine pouches in this species. The embryos are a trifle 
larger than in P. y fetllax and the second egg membrane ^5 also larger 
in P. d/a bius. In other rejects thes^species are remerkably alike. 
Without staining specimens of both species and comparing them care- 
fully they would be declQjred to be identical. 

This species may be differentiated from P. >| 3 gle e t us 
bg the smaller size of its proglottids and by the larger size of its 
embroyoj , The proportions of the cirrus-pouches are also very 
different • 

P. d ub ius is much unlike P. f> ->rcae and P . c 3rnuae _ in 
size, size of jueker^in number^size anc arrangement of tes^s and in 
length of cirru; -pouch. 

P. d ub ius differs from P. esociSjP. Agon is and 

P. ^sterostei in size and in the presence of a fifth suckers 
There is also considerable difference in the length of the tfirrus- 


1884: Taenia 

1884: Taenia 

1892: Taenia 
1892: Taenia 


(Pl.2figs.33 ~3») 

ocellatus hokke, 1884: 13-14, in part? 

Zschokke, 1884-14-15 

Kraemer, 1892530-55 

Kraemer, 1892^72-76 

ITufer, 1905 in part? 

Fufer, 1905 in part? 

f ilicollis 

ooellat a 

1905: Proteocephalus ocellata 
1905: Proteocephalus lcngic<e>ll is 

SPECIFIC DIAGNOSIS.- Characters of the genus. Cestodes 
quite small . Length up to 100 mm. Max. "breadth up to 1.20 mm. 

Head somewhat globose, 0.150-0.200 mm. in breadth by 
0.105-0.150 mm thick. Head well set off from neck. Suckers situ- 
ated near "broadest zone of head. Dietmeter of suckers 0.064-0.085 
mm. Diameter of sucker opening about 0.040 mm. Fifth sucker 
present, functional, about 0.058 mm. in diameter. Feck narrow, 
length 2-6 mm. First proglottid© quadrate or longer than broad. 
Mature proglottids quadrate or broader than long, 0.34-0.46 mm. 
broad x 0.34 mm. long. Ripe proglottids longer than broad, rarely- 
measuring as much as 1.19 mm. broad x 1.36 mm. long. Segmentation 
not very distinct, margins smooth. 

GENITAL ORGANS.- Genital sinus marginal, alternating, 
situated near middle of the proglottid. 

Male organ s.- Testes 30-35 in number, irregularly scatter- 
ed in single leayer between vitellaria and anterior to ovary. 
Diameter of testes 0.037-0.060 mm. Cirrus-pouch in mature proglo- 


ttids 0.196-0. 255mlong in ripe prcglottids. Ratio of length of 
cirrus-pouch tc breadth, of proglottid 1:3-1:2. Cirrus and ductus 
ejaculations nearly straight. Length of protruded cirrus 0.026 mm. 
Vas deferens forming a mass of coils in maddle of proglottid. 

Feznal e org ans .- Typical of genus. Vaginal opening 
anterior to ci rrus-pcuch . Vaginal sphincter weak. 

Ovary hilohed, lobes solid, thick, elongated ovoidal in 
form. In ripe proglottid ovary lies in trianglar area between 
pouches of uterus. 

Vitellaria lateral, and sparse. Organs of interovarial 
space as in genus. Uterus ventral, in ripe proglottids with 6-8 
lateral pouches on either side. Uterine pores 2-3 in number. 

ECtGS.- Outer shell of uterine eggs thin arw. hyaline, 
middle shell granular 0.036-0.041 mm, embryo 0.031-0.0336 mm. in 

HABITAT.- Intestine of Coregonus f era, Lake Lu*e<gTir?e 
Switzerl and. 

TYPE.- He. 09.9 in Prof. H. B. Ward's collection. Slides 
from same lot in Professor Ward's collection. 

HISTORICAL SUMMARY.- Zschokke (1884) reported and desc- 
ribed in a very gener- 1 way five species of Prot eocephalus . His 
Taenia salmcnig. umblae and Taenia t orulosa are discussed in their 
proper places. The other three species, his Taenia oflellata , T. 
f ilico ilis t and T. longi c ollis can best be discussed together and 
in this connection. The small table shows the distribution of the 




five species in the hosts examined by him from Lake Geneva. 

Host s 

f era 

Salmo Trutto : Perca 
umbla variabilis fluvia- 



vul garia 


Taenia frT. longi- T.long i- 


longi - 

T. ocell 
at a 

Prot eo- 
cephalus : 


t orulosa 



salmon l s : 









f ilicollis 

t omlosa 

t cpil oa 

The Taenia fj lico .l l Is of Zschokke?s description (1884:16- 
17) is from Perca fluviatilis and may be the same as the specimens 
from Perca fluvi atilis received by Dr. H. B. Ward from Professor 
Parona which I have named P. dub ins n.sp. It is certain that 
Zschakfcas description and drawings of his specimens from Perca 
fluviatilis do net agree with Schneider's P. percae (0 .P .Mu Uler ) . 



The specimens cf T. oc ellata which Zschokke reported from Perca 

f Haviatilis are probably only longer specimens of his T. f ilicollis . 

The specimens reported by Zschekke from Esox lucii>s were 
probably Prot eocephalus e socis Schnaider. It is doubtful if Lota 
vulgaris harbors the same species of parasite as do the Salmonoid 
fishes. Zschokk^s report on this point needs confirmation. 

The Taenia ion ico n is and T_. oc ell at a from Coregonus fera 
are probably identical. ZschokkS ' Rescript ion of his Taenia longi- 
call is was no doubt based on specimens from Coregonus fera for he 
wrote: "Pendant me recherches je lai trouve en grande quant ite dans 
les appendices pylori ques et dans les intestines greles de Coregonu s 
fera , et une fcis, en Fevrier, enkyste sons sa forme larvaire dano 
le foie de Salmo Umbla. " 

Fortunately Dr. H. B. Ward has received seme specimens frorr 
Professor Zschokke labelled, " Taena ccell ata , Coregonus fera ." This 
has been found to be identical with Kraemer's Taeni a ocellata from 
Coregonu s fer a, Lake Lucerne. This proved to be a new species to 
which I give the name of P. f allay. . It is highly probable that 
the specimens from Ccregonu s fera. identified byZSchokke (1884) as 
longicall i s belong to this sane species. His description of his 3c. 
longicol] is differs from that of Kraemer (1892) chiefly in length. 
The small head, large cirrus-pouch, the proportions of the proglott- 
ids, position of the genital pore; size of the cirrus-pouch agree 
pretty well with the facts for P. fall ax n.sp. 

Zschokke (1864) presented no data to show that he had made 


a comparative study of the specimens collected by him frcrr. Salm o 
Umbl a and from T rutta variabilis . It is scarcely possible to de- 
termine where these specimens belong. and speculation without seme 
basis of fact is worth but little. 

taken from Ccrercnus f era , Lake Li ccrme in July identified these 
specimens as Taenia f ilicallis Rud. Later in the same season (about 
Septe. 1) he took more specimens from C oregonus fera. These he 
identified as Taenia ocellat a on account of their greater length. 
His specimens collected in July measured 50-60 mm. Those taken by 
him in September measured up to 100 mm. Kraemer now made a careful 
comparison of th^se two forms . He found that they agreed in every 
particular as to size of the head, of the suckers and as to the 
presence of a fifth suc v er. In the internal morphology of the 
proglottids ard in the histological structure of the organs the 
two forms also agreed perfectly. Moreover among his specimens he 
found gradations in length from the shortest to the longest. On 
these grounds Kraemer concluded that Taenia f ilicoilis Rud. and 

Taenia ocell a ta R ud. were one and the sa-r.e; that in fact the smaller 
Taenia f ilico "I is was but an ircompletly developed Taenia, ocellata. 

smaller specimens in his possession belonged to the same species. 
However, his conclusion has nothing what soever to do with Rudolphis 
Taenia ocellata and T. f ill i call isjror tht very good reason that 

Kraemer (1892) as 

a result of his study of specimens 

Kraemer was correct in concluding that these larger and 

Kraemer' s specimens belonged to neither of Rudolphis species. 


Fraemer in his papei^gave no adequate reasons for con- 
sidering- that the Taenia f ilicQlli s and Taenia ocellala of Rudo/phi 
were identical. His specimens were neither frcm Rudophis type host 
nor from his type locality but from a very different typehost and 
type locality. Zscbekke had material frcm Perca f luviat ili s yet he 
described what he condidered to be Taenia f ilicQllis rather than 
Taen ia c eel lata from that host. Neither Zschokke nor Kraemer had 
material from Gast erosteus . 

Kraemer' s specimens beyond a doubt belong to the species 
P. f allax n.sp. to be described. 

Riggenbach (1896) accepted /(raemer's conclusion concerning 
the identity of Taenia o ce ll ate and T aenia f ili co llis . He ap- 
parently made no comparative study of the forms. 

From North American fish Benedict (1910) described a 
species which he considered identical with Kramer's Taenia, ccellat a 
and po made it Pre t eccephalus ccellata (Rud ) . That, however, was 
a misdeterminat ion for Benedict's specimens have been shown to be 
a new species (see P ." exiguus n.sp.) 

Kufer (1905) may have been dealing in part with this 
species in his statements regarding P. oc ell at us a nd F,l ongicfl,llis 
from Coregonus f era. 

MATERIAL.- The material on which this species is based 

bears the label "T . c cellata, Cosegonus f era." It was secured from 
2rof, Fritz Zschokkeby 

Prof. H. B. Ward. It now bears the number 09.9 in Professor Ward's 
collection. Some of this material was stained in haematoxylin and 


mounted in "balsam. Other specimens were cleared in glycerine. 

Comparison of material with Kraemer 's Rescript ion : - 
Study of tbese preparations made evident the fact that this species 
and Kraemer's Taenia ccellat a were morphcl ogically identical. 
The fact that Kraemer's specimens were also from Coregonius f era 
and that doubtless they came from the same locality was one of thfe 
considerations which caused t v is identity tc "be suspected. On 
account of this identity Kraemer* s description will he used as a 
source of comparative data and all of Kraemer's data on his T. 
ocellata v/ill be grouped tcgether. 

EXT EE IT AL CHARACTERS . - The larger worms at my disposal 
were unfortunately "broken into pieces. One complete strabila 
measured 27 mm. long with a max. breadth of 0.459 mm. A small 
piece in the same lot had a max. breadth of 1.20 mm. Kraemer reported 
specimens of T_. ftc ellatus 30-60-100 mm long. 

He reported a. breadth of 0.114 mm. for the head and 
0.038 mm for suckers. The last measurement must be that of the 
sucker opening and not the max. diameter of the sucker. 


My measurements in mm. of seven scolices and suckers 



: "breadth of head 

: length 

: of 

: head : 

: suckers max. 
: diameter 

: fifth 
: sucker 


: 0.187 mm. 

: short : 

0.085 : 

not observed 


: 0.185 mm. : 


: 0.080 : 

n h 


: 0.170 

> II 

: 0.064x0.064 : 

ii ii 

4 ! 

: 0.185' 

. II 

: 0.068 

; n it 


: 0.204 ; 


: 0.082x0.074 

. n it 


: 0.153 byU105 thick, 


• •• it 


: 0.187 by 0.153 " 

> II 

: 0.079 

: 0.058 mm. 

The opening of the suckers in No. 4 measured 0.042 mm. 

There is here some discrepancy between "Kraemer's figures 
and my own. This may be due to an error in manipulation, to the 
fact that Kraemer happened to measure a very slender head, or that 
he happened to measure it in the dorsoventral (the shorter) dim- 
ension. As will be noted the diameter of the sucker as given by 
Kraemer is about the size which I give for the sucker opening. 

The head^is somewhat globular in shape. The suckers are 
situated at or just above the broadest zone and they are directed 


somewhat anteriad. In my material the head is readily distinguished 
from the thin neck. 

The neck is 0.076 mm. broad (Kraemer ) --0 . 136 mm. "broad 
in my measurements. Kraemer states that it is long but he points 
out that in stained preparations part of that which appeared to be 
neck was in reality made up of young proglottids. In my glycerine 
mounts the neck sometimes appeared to be up to 6 mm. long while in 
stained preparation 2.38 mm. was the maximum length. 

The first proglottids are about quadrate or they may be 
longer than broad. Kraemer gives the following breadth for the 
various proglottids, first long proglottids 0.228 mm. broad, the 
quadrate anterior proglottid 0.532 mm, the sexually ripe mid- 
proglottids scarcely 2.0 mm, the ripest end-prcglott ids up to 2.0mm. 
My measurements of similar proglottids are as follows: The first 
proglottids 0.136 mm. broad x 0.17 x 0.30 long, quadrat e mature 
proglottids 0.34-0.46 mm. broad x 0.34 mm long, the sexually ripe 
proglottids, 1.19 mm bread, 1.36 mm. long (only four proglottids of 
this size were observed.) 

The segmentation of the w era is not very distinct. The 
margins are quite smooth except for very slight indentations at the 
junction of the segments. The segments are closely joined together. 
Tbe genital pore is not marked by a papilla. 

The common genital sinus is marginal, and is situated near 
or slightly anterior to the middle of the segment. The position of 
the sinus alternates irregularly in the chain* , 


INTERNAL ANATOMY. - Kraener worked cut in ccnsderable 
detail the structure cf the cuticula, the musculature, the paren- 
chyma, the nervous system, the excretory system, and the histological 
structure of the various internal organs. In the main I shall not 
discuss these points, however, some of Kraemer*s conclusions are 
incorrect. These errors I shall indicate when possible. 

GENITAL ORGANS.- Male Org^s .- In may preparations the 
testes number 30-35 and they measure 0.037x0.053 mm. Kraener 
recorded 27-30 testes a diameter cf C.057 mm. The testes^ lie in 
a single layer in the field bounded by the ovary, the anterior 
margin of the proglottid and the vitellaria. 

The c irrus-pouch^ is 0.196-0.255 mm. long, in fully mature 
proglottids, where its ratio to the proglottid bre&di is 1/3-1/2. 
In fully ripe proglottids it is more or less concealed by the 
uterine pouches. In a few ripe proglottids in which it was net 
contracted the cirrus-pouch measured 0.370-0.420 mm. in length. 
Apparently in his speaies the cirrus-pouch becomes larger after 
sexual maturity is attained. The cirrus-pouch is slender. Its 
inner end is slightly curved upward toward the dorsal surface where 
its is attached to the walls of the derm ■■ muscular sac by strong 
muscle fibers. This inner end is usually somewhat swollen making a 
little vesicle. Within this vesicle one or two sinuous curves of 
the ductus ejaculations may be seen. Adjacent to the v'escile is 
the constricted region of the cirrus-pouch. Here even in toto 
preparations strong circular muscle fibers may be seen. In this 


region the ductus passes over into the straight slender cirrus. 
Just beyond this constricted region the cirrus-pouch again dilates 
a little and here the circular muscle fibers are not prominently 
developed. From a cirrus-pouch so constricted the cirrus itself 
is usually protruded. The length of the incompletely protruded 
cirrus is about 0.026. Xraemer states that the cirrus protrudes 
2-3 mm. This must be a typographical error for a cirrus as long 
as 2-3 mm. would require first a very large cirrus-pouch and second 
many coils of vas deferens within the pouch. 

Kraemer's drawings and data do not lead me to believe 
this is the case. His firures shows a protended cirrus probably not 
over 0.3 mm. in length. The vas deferens forms a mass of coils at 
the inner end of the cirrus-r.ouch. In mature proglottids this 
knot lies in the middle field, even reaching past the middle. 

Kraemer undoubtedly made some errors in his description 
of the cirrus of this species. Benedict (1900) pointed out these 
errors thus: 

Kraemer's description is widely different. His drawing 
shows a slender tube with hooks, representing the cirrus, and 
followed by a proglott i s-like arrangement, and then several coils of 
the vas deferens within the pouch. The drawing has an unreal ap- 
pearance on the first glance. The cirrus, with its curved hooks, 
is imbedded in the tissue, which must be torn through before it 
could be protruded. The proglottis-like appearance of a portion of 
t>ie cir>ns, as he drew it, was undoubtedly due to the way in which 


the circular muscles were cut in sectioning. He drew the cavity 
of the cirrus, into which the cut ends of the circular muscles 
projected, as the external outline of the cirrus. The coils would 
"be necessary, according to his theory, but are not to be found. He 
says that the muscles, which he calls "the roots," are for re- 
tracting the pouch, which is sometimes throst outwards for some 
distance through the opening. This protrusion is really due to an 
pagination of the free distal end of the inner tube. 

Fe-nale Organs .- These in arrangement are typical of the 
genus. The vagina^ opens always ant eri ctivt o cirrus-pouch. Near 
its opening at a distance of 0.041 mm. according to Kraemer, it 
possesses a weak sphincter muscle 0.026 mm. long by 0.034 mm. broad 
over all. In its course to the interovarial space it crosses the 
bulbous end of the cirrus-pouch or extends beyond the end of the 
pouch before bending I have not noted in toto prepara- 
tions the dilatation in the middle portion of the vagina as desc- 
ribed by Kraemer There are no coils of the vagina anterior to the 

The ovary is bilobed, the lobes connected by a slender 
midpiece. The lobes are solid, fairly thick bodies of elongated 
ovoidal shape. In mature proglottids the span of the ovary is 
about 0.350 mm. In ripe proglottids the ovary in stead of becoming 
"iore elongated and flattened against the posterior wall of the 
segment shrinks in size and the two lobes become quite closely 
pressed together in a s-nall triangular space bounded by the posterior 



proglottid margin and by the walls of the uterine pouches. This 
triangular space formed "by uterine pouches in the posterior end of 
the proglottid is a very characteristic feature in this species. 

The vitellaria are elongated follicular glands in the 
lateral margins of the proglottid. The vitellaria are sparse. 

The organs of the int er-ovarial space are probably as 
Benedict has described them for his species and not entirely as 
Kraener has described them. I have not made sections of this form 
so I have been unable to trace the connections of the ducts. 

Kraemer established the presence of an ootype, an oocapt 
and a shell-gland. The paired vitelline ducts do not originate so 
far anterior as Kraemer has figured them: They probably unite to 
form a single vitelline duct as in other Proteocephalid species 
where this has been investigated, 

Kraemer noted in toto preparations that the vagina anterior 
to the ovary was apparently divided, one branch going to the ovary. 
The second tube is probably the uterine passage which can frequently 
be seen in toto preparations and which gives rise to some trouble 
unless its function be understood. 

The uterus^is a median tube in mature proglcttids. In 
ripe proglottids there are 6-8 lateral pouches, on either side of 
the median tube. These lie in the ventral field and come to occupy 
nearly the whole proglottid. By the densly p^cfted uterus the 
vitellaria are pushed farther toward the margins, the testes close 
up to the dorsal surface and the shrunken ovaries are crowded into 


a small triangular space in the posterior part of the segment. The 
ripe proglottid is practically a thin walled sac divided up into 
compartments by the thin septa of the uterine pouches. 

The eggs are discharged thru 2-3 uterine pores or thru 
a rift down the ventral surface which is caused by a further split- 
ting of these uterine pores. The usual number of uterine pores 
is 2 or 3 but occasionally a proglottid is found with a single 
pore. Kraemer describes a single uterine pore in the middle of 
the segment. The eg^s have shell with three membranes, the outer 
hyaline, variable in size, a middle heavier and more granular membr- 
ane, and an inner thin membrane a enclosing the embryo. The embryo 
measures 0.031-0.0336 mm. in diameter. Such a wide variation is 
probably due to the different forms which the embryos assume. Some 
are spherical others cvaridal. The middle membrane measures 0.036- 
0.041 mm. These measurements show very well the limits of variation. 
The outer membrane was not measured. 

RELATIONS.- This species is readily differentiated from 
P. perca e by its smaller head and suckers. more slender neck, and 
smaller proglottids. it is less robust than P. p ercae . It differs 
from P/ p ercae in having a much shorter ci rrus- pouch, fewer testes, 
a shorter ovary, and a larger embryo. 

It is readily distinguished fromP. t oml c su s by its smaller 
size and by its lack of a fifth sucker. It differs from the P. 
1 ongicoll i b Rud. as described by Linstow in having a smaller head 
with much smaller suckers. In the P. longicall is of Q. Linstow the 


testes are much larger than in P. fall ax and they are in two (?) 
fields. The cirrus pouch is shorter in P. l ongicallis . The 
lateral pouches of the uterus are fewer in number and the embryos 
are smaller than in P. fall o>. rep. 

P. f all ax n. sp. varies from the P. cernuae from Acer^0a 
cernua in being more slender, with a smaller head and less muscular 
suckers. The latter species has broa.der but narrower proglottids, 
a shorter cirrus-pouch, more numerous and much larger testes, a 
longer span of ovary, and a smaller embryo. 


P. fall ox differs from P. g- aotoroot oi in being longer, 
in having a fifth sucker, a longer cirrus pouch, fewer testes, and 
larger embryos. 

P. fall ox much resembles P. exig uus but it differs from 
that species in having a larger head, larger suckers, larger fifth- 
sucker, a. longer neck, more numberous a,nd larger proglottids, fewer 
testes, a shorter cirrus-pouch, fewer uterine pouches. The embyos 
are larger than in P. exigu.u s . 

This species further resembles P. pusillus Ward. It 
differs from that species in the character of the segmentation, in 
the smaller size of its suckers, in the position of the genital 
aperture, in the smaller number and size of testes, in its longer 
cirrus-pouch and in its fewer uterine pouches. 

P. fall ox resembles P. ag^onis Barbieri but it differs 
from that species in being somewhat larger, in having a fifth sucker 
fewer testes, and more uterine pouches. 


than P. f allax 

P_. esocjL g G. Schneider is smaller . 1 n s] and it lacks 
a fifth sucker. It moreover has a larger and. more numerous testes 
than P. fallEUBp. 

P. fall ax most closely resembles P. dubiu s n.sp,^. Perca 
f luviatit i s hut differs from that species in having a much smaller 
number cf testes, smaller testes, a relatively shorter cirrus-pouch, 
fewer uterine pouches, a slightly smaller embryo, and a small second 

membrane. In size, in size of head, size of suckers, they re- 
semble each other remarkably. 


(PI. figs. ^-J2.) 

1780: Taenia e^perlani Acharius, 1780 :b2. 

I78y: Taenia salmonis V/artmanni Froelich, 1789:24. 

1790: Taenia Froelichii Gmelin, 1790; 3080. 

18U1: Taenia longicollis Rudolphi, 18U1-.11S. 

18U0; Alyselminthus l ongicollis Zeder, 1800; 258. 

1803; Halysis longicollis Zeder, 1802; 333. 

1810: Taenia longicollis Rudolphi, 18iO;v2:io7. 

1819: Taenia longieollis Rudolphi, 1819:149. 

184b: Taen i a longieollis Dujardin, 1845:58b. 

1850: Taenia longieollis Mesing, 18bO:bl2-bl3. 

1891: Taenia longieo llis V. Linstow, 1891:565-576. 

1894: I chthyotaenia l ongieollis Lonnberg, 1894:803. 

SPECIFIC DIAGNOSIS.- Characters of genus. Cestodes of 
medium size. Observed length (Yon Lins tow's) 46.4 mm. by max breadth 
ol" 0.99 mm. Length 20-200 mm. and breadth 1-2-2.25 mm. (Dujardin). 
Scolex 0.43 mm. broad. Keck ©.3 mm. broad x 3.2 mm. long. Suckers 
circular , 0.12-0.14-0.18-U.19 mm. in diameter. Fifth sucker half 
as large as others* First proglottids 0.45 mm. long x 0.71 mm. 
broad. Proglottids at and of first third 0.b3 mm. long x 0.99 mm. 
broad, posterior proglottids l.oa mm. long x 0.85 mm. broad. End 
proglottid triangular. 


EXCRETORY SYSTEM: - One pair large Central vessels* 
Three paris small dorsal vessels. Many anastomoses oetween vessels, 
-.osterior transverse commissure oetween ventral vessels. Circular 
commissure posterior to suckers. All excretory vessels much contorte :. 

GENITAL ORGANS.- Genital pore irregularly alternating 
situated at end of anterior, one i'ourtn oi' lateral margin oi 

Hale organs.- Testes spneroidal, u.09 mm. in diameter, 
2b (?) in each segment, arranged in twcP^fields near vitellaria. 
Vas dei'erens a close mass in median j ield. Cirrus-pouch spindle 
shaped, muscular ,0*11U mm. broad, extending just thru vitellaria. 
Cirrus clubshaped reaching u.2 mm. past margin of segment. 

Female organs.- Ovary biiobed, posterior. Lobes slender. 
Vitellaria lateral, follicular. Uterus with three lateral pouches 
Vagina anterior to cirrus-pouch not crossing cirrus pouch. Sphincter 
Vaginae? Receptacuium seminis? 

EGGS.- Unripe eggs U.ui66-0.0iy6 mm. in diameter. 


Habitat: In intestine. 


Coreg-onus lavaretus 
Coregonus maraenulr 


Lake Lucerne 

Lake Geneva 

Rossiter £• 
liicolaikon K. 

C. schinzii nelvetic iff Lake Lucerne 

Lake Lucerne 

Coregonus T/artmanni 

q C. term 
C. fei 
C. alt aia 


' £• exiguus albeilus 
ostierus eperianus 
usmerus eperianus 

Osmerus eperianus 

Trutta trufca 

Trutta trutta 

Trutta lario 

Salmo thymoiius 
S almo a lpinus 
Salmo salvelinus 

Salrao saivelines 

Salino umbla 



Museum Vienna 

Lake Lucerne 
Lake Geneva 







liMt er 


U. L ins tow 




Mart in 



Rudolphi, iaiu,v2 

Rudolphi 1810, v2: 

Rudolphi iyiu.v2: 


Uuier iyob:7b. 

Zschokke 1084:11- 


Mum ing,iyy«:36. 

Hui'er l ( JUb:7b 

Bufer iyub:7b. 
Kudoipni Idiu.v2: 
Kudoipni l8lC,^ 

v2« 107. 
U.Linstow 18yi:5b 


Rudolphi 1810, V2: 

Rudolphi 1810 t Y2: 

Kudoipni iaio,V2 

Kudoipni 1810 , V2: 

Rudolphi 1810, V2: 
10 7. 

U. Linstow 1878: 

Wufer lyob :7b. 

Zsehojfcke 1884: n 



V. Katz : 

; von Katz, iuy7:it»y. 

? Esox iucius 

: Lake 


: rful'er : 

liul'er ±'JOb:7b 

? ttsox UC1US 

< .ferca lluviatilis 




Huier iyub:7b. 

1 Alburnus lucidus 


Lucerne ; 


imrer iyub:7b. 

4 Squaiius cepftaius 




jaufer iyub:7b. 

? Squalius l evciscu s Lake Lucerne Huler Nufer iyub:7b. 



HISTORICAL SUMBAHX.- Kudolphi 1181U, v2: 1V7) gave a 
diagram and description o±" Taenia lcngioollis w hich is here quoted 

l *TAKHIA LUliGIuuLiils K.- Taenia : c apite truncato . c oiio 
longissimq . articuii s subquadrati s. ovariis racemosis. 

Froiich im naturi'orsch. 24. p. 124. Tab. 4. f ig. 20.21. TaentA. 
Salmonis v/artmanni. 

Gael. Syst. nat. p. 3l)tfu.n.yi. Taenia i'roelichii. 

Zeder Nachtrag p.2b8.Alyselminthus longicollis. 

Kudolphi in '.vied. Arch. 111. 1. p., 113. Taenia longic. 

Zedor Uaturg. p. 333. n. 9. Halysis longic. 
Hab. in intestinis Saimonum. In Saimonibus v/artmanni 
plurimis ifroelichius Augusto copiosam; in s. Trutta zeaerus; ego in 
s. lavcrot o Ha jo ot uaraenulis plurimis Februario, reporimus. In 
Salmone Eperlano a se inventas am. Treviranus mecum communicavit . 

Descr. vermes unum ad septum oollices longi, lineam di- 
midiam vel integram lati. 

Caput depressura, truncatum, exiguum; osculis orbicularibus , 
binis tarn superioribus , quam inferior! bus , capitis rnargini antico 
a oproximatis , ut sub ejusder.i raotu saepe antica oraniague simul 
appareant. Collum cum capite continuum, iongissiraum, tertiam circiter 
totius longitudinis partem Bibi vindicanc, deprossun, margine obtuso 
integerrimo. Corpus depressum, articalis anterioribus brevissirais , 
reliouis subquadratis, marginibus singulorum antico et posico rectis, 


lateralCo'ous rotundatis, ut totius vermis latera crenata appareant. 
Articulu ultimus obtusus. Ovaria in articulis posticis, inde vel 
caerulescentibus , vei nigrescent ibus, distincta, raoemosa sive 

Obs. 1. An Scolex tetrastoraus supra dictus hujus Taeniae., 

Taenia iongicoiiis R. (Cont.) proles nondum articulata? 
Sed reliquarum Taeniarum foetus semper articulatos vidi, neque caput, 
pars postica acuta conveniunt. 

Obs. 2. Hujus speciei cum insequeatibus, ob articulos 
margine postico vix incurabentes, ob capitis coliique formam, 
affinitas magna j differentiae tamen specif icae singuiarum discrimen 
satis indicant. 

Obs. 3. An quas Stellerus (Pallas II. Nord. Beytr.l.l.p, 
1QZ) in Eperlani vesica natatoria reperit, Taeniae hue pertinent? 
Conf. n.113. 

Rudojpiii (1819:149) quoted the diagnosis which he had 
used before (Rud. iblU. V2:1U7 ) . He stated that he had found the 
species in Coregonus iavaretus ax Greiiswald, in maraenula 
and in Osmefrtis eperlanus at Berlin* Froeiich iouna io in 
Coregnuus Wartmanni . Frdlicn and Zeder i n Trutta trutta . It was 
found in S . Thymallus» Thymailus vexiilifer in the museum at Vienna. 
Harxin found ix in Saimo alpmus and in T rutta farie . 

Dujardin (ia4£>:abo) ad L ed nothing to the data of the 
e .-lier authors. He staxes;"Ii (Rudoipnij, en outre, que f* Taenia 
r enkina de Schramm est identiq..e avec celucici, parce que le 


Salmo uartmanni est nomme renicen dano L'Allemagne mendionaie" 

I nave not Deen able to justify this statement. 

jjiesing ( i8bu:t>i;c:-i3j gave nothing new. U. Linstow 
±«YO:^b^) in his catalogue or Bfctozoa gave Salmo saivelmus as a 
nost 01 this species. 

zachokke ( 1««4:11, 14-16 j identified and described some 
specimens from Ooregonus f era and Saiiso umbia 1'rom Lake Geneva as 
T. longiconis . A careful comparison ox this description with 
his description of Taenia oceilata likewise reported by him from 
these two host£ .s well as from several other hosts of the same 
locality compel me to believe that his *V» o cellata and his T. 
l ongicoilis may have been identical at least m part. The chief 
difference in the description is as to the length of the two forms. 
In his description of certain organs in the two forms he used 
almost identical phrases. It is my opinion that Zschokke depended 
upon a single character, viz. length, to distinguich the two 
forms. Kraemer (ltf'J2J one of Zschol&e's students evidently did 
the ^ame in his study of P. f all ax (T. oceilata ) • If Zschokke 
did as I suspect, tnen he may have had to do with a single species • 
In such a case his description for the two forms would necessarily 
be much alike. 

Zschokke' s description of T aenia longicoilis does not 
agree wit the description and figures or that species. by Von 
Linstow ^yy/:bbb-67b) Yon Linstow' s specimens were taken from 


Coregonu s eperlanu s. one of the hosts in which Rudolph! round 
T. l ongicollis . It is therefore probable that U. Linstow's 
specimens rather than Zscho^:es belong to Rudoiphi's species. 
Moreover, von J, in tow's description agrees more completely with that 
of Rudoiphi's tnan does Zschofcke's. 

Lonnberg (l8y4:U0s) included Taenia longicol Lis Rud. 
in his list of species of his genus Icnthyotaenia. 

Zsohokke ( 18 v J6:77^-777) listed Icnthyotaenia longi c ollis 
only in Trutta iarip from Lake Geneva, Thanks to Prof. H. i3. '.Yard. 
I nave been able to examine parts of a strobiai oi Zschokke's 
"T. fronglcoliis aus Porelle" ( Trutta far is ) and I am able to state 
positive by tnat it is not the some species as V, Linstows T. 
longicollis . ITor is it the same as the specimens from Coregonus 
fera which Zscnoicke sent to Professor Wfcrd as T. ocellata (No. 
0C9^in Prof. H. 3. Ware's collection anion is the type of P._ fall ax 
. Sp. ) 

V. Ratz (iy v J7:i&yj listed Ichthyotaenia longicollis 
from Esox luciu sfrom Lake Balaton in Hungary. This is probably 
a ratsdetermination. v. Ratz gave no description ao not evan a 
: probable determination can be made. 

- Muniing (iay«:bbj reported Ichthyotaenia longicollis 

i (RudJ 



rrom Coregonus albuia . at ^icolaiken and from Qsmerus eperlanus 
at Rossitei) in East Prussia. He gave no description. 

Nuier (iyOD:7oj in an investigation ol" Lake Lucerne 
reported Proteocephalus l ongioollis from Perca i'luiatili s. Alburnus 
lucidus . SgualUls cepnalus . S. Leucisous . Esox lucius Coregonus 
7/artmanni nobilis . C. e xigqus albellus .C. schinzi i helveticus 
and Salrao salve linus . It is extremely doubtful it" tho specimens 
ol Proteocepnaius lound by Hufer in oiher than tne Sairaonoid fish 
belong to the species P. longicollis . llufer's tabulated description 
( Nui'er iyut>*147j is almost entirely based on the work of Von 
Linstow (itfyij. The character oi lufer's work has been discussed 
at some length in connection with P. macrocephalu s^ page , and 

P. t orKlosus page , Many ol" the statements made in those places 

regarding his work apply here. 

SOURCE OF DATA. - Tne description is based on the 
paper by Von Linstow (10y±j. His material was collected from 
Qsmerus e perlanus . one of the hosts in which Kudolphi found Taeni a 
longicollis . 

EXTERNAL CHARACTERS.- Von Linstow 1 s largest specimen 
! measured 4b .4 ram. long with a max. oreaatn oi u.'jy mm. At tho 
posterior end the worm was 0.83 mm. broad. The scolex was 0.43 mm. 
broad, followed by a neck 0.3mm. broad ana 3.2 mm. long. The 
first progiotLids were 0.43 m . long oy o.Vi mm. broad. Tnose at 
the end or the first one-tnird oi the etroona where the cirrus 
v/ere developed were o.t>3 mm. long by O.y'J mm broaa. Tnen the 


length 01 tne progiottids increased until tne last proglottid^ 
were 1.U3 mm. long ana u.««5 nun. broad, Tne end-progiottid v/as 
triangular and drawr. out posteriorly. Tne specimen was incompletely 

The i'our suckers were circular in outline, 0.12-0.14- 
0.18-0. iy in diameter. A fifth sucxer at the apex oi' the scolex 
measured one-half the diameter oi the others. 

EXCRETORY SYSTEM.- The excretory system is made up oi 
two large lateral vessels, one on either side just outside of the 
vitellaria, and six smaller vessels, three on either side lying 
dorsal (Jp tne larger vessel. All the excretory vessels are greatly 
twisted a. id many anastomoses exist between them. In each 
proglottid about u.utfu mm. anterior to the posterior margin is a 
transverse commissure connecting the pair of ventral excretory 
vessels. The two larger vessels measure u.uiy mm. broad and the 
smaller vessels about U.UU78 mm. In the region just posterior to 
the suckers, a circular commissure 01 excretory vessels is formed 
from wn'ch branches extend anteriorly into the head. All the main 
vessels end posteriory in a small vesicle in the end of the end- 



GE1TITAL OK G AN 3.- The sex opening is marginal^ irregularly 
alt rnaxing rignt and left and is situated in V. Linstow's drawing 
at the end of the anterior one-fourth of the proglottid. 

( pH./wJ 

Male organs The testes are large, s'oneroidai, measur 


ing up to u.u mm. in diameter. They are about 25^tests in each 


se^monx. These ere situated in two fields near the vitellaria. 


In a drawm~ or a transverse section Von Linstow figures the tests 


: as occunng two layers. This causes rne to suspect that the total 
number of test's (2t>J as stated by Von Lmstow is too small for in 


none of the soecies of P roteocephalu s thus for examined by me have 
the testes been arranged in t. o layers except when the number of 
testes is about rifty or more. Two layers oi testes is not always 
the rule in those cases. Moreover, in that single transverse section 
| ten testes are figured^a larger mumber than one would expect to find 
in a transverse section of alorgeproglottid which has only tib testes • 
:3ince the proglottid which Von Linstow delineates is ripe, it is 
probable that he overlooked the testes or the median region where 
the eggs would be most densely packed. In the transverse section 
to which rel'erence has been made the region dorsal to the coils of 
vas deferens is 1'ree rrom testes, a condition which exists in all 
the other species or the genus whether the number or the testes be 
few or many. If Von Linstow's statement be Correct that the testes 
are in two lateral fields one on either side of a free median zone 
then this species is the only exception to the rule that the testes 
in this genus are irregularly scattered between the xatoral 
: vit^e llaria. Ail these considerations cause me to douot the 
j accuracy or Von Linstow' s statements regarding the number and 
position oi the testes. 


The vas deferens forms a thick mass oi' coils in "che 
middle oi" the proglottid. This mass apparently does not extend 
over "of sne cirrus-pouch. The cirrus-poucn is spindie-snaped, mus- 
cular, mm. broad. In Von Linstow's figure the cirrus-pouch 
extends but little witnin the viteiiaria. 

The cirrus in the 46#4 mm. specimen was visible 15.8 mm. 
posterior to the head. It is short and ciut>ehaped, mm. long, 

and extends U.£ mm. beyond tne proglottids margin. 

Fema le organs . - The biiooed ovary^iies in the posterior 
region oi the proglottid. The viteiiaria lie right and left in 
the lateral fields of the v-oglottid. They discharge the yolK - 
ceils tnru paired vitelline ducts. The<j i ducts pass to the 
interovariai space wnere they unite to form a common duct which 
empties into the ootype. Thus r<*r V. Lindtow ib correct in his 
description of tne organs of the interovariai space. According to 
him the vagina discnarges into the obtype as do also the two lobjs 
of the ovary. He show no visible means for the escape of the 
fertilized egg from the obtype. Von Linstow nas probably mistaken 
tne oocapt for theotftype. Honticeilil iy.;i :16E) in discussing Von 
Linstow* s description or T. longicolii s arrived at almost the same 
conclusions regarding tne ob'type and oocapt in that species as I 
have stated nere. Ha nas erroneously xigured the vitelline duct 
v/hlch pa ses near the oo'capt and the shell glads lying near, as 
discharging into it. 


The barrel-like form oi" the ootype as delineated by him is much 

\ more typical oi x;ne oocapt tnan of the ootype v;hicn is much less 

conspicuous. The vagina or Von linstow's figure 4 is doubtless the 

oviduct. The organs ol" the interovarial space doubtless stand 

in the same general relations to each other in this species as 

they have been round in other members or the genus. 

The vagina opens into the common genital sinus, always 

anterior to the cirrus-poucn. Without crossing the cirrus-pouch it 

extends in a curved course to the interovarial^ pp0h^ where it 

lorms several coils . Von Linstow did not mention the presence oi' 

a shincter vaginae. 

The uterus has three large lateral pouches on either 
side. A uterine pore is lacking. 

2GGS.- The unripe eggs measure u.ulbb-U.'Jivb mm. in 


DEVEL0PL1EM] . - The plerocercoid of this species is 
found in the liver of the same host. 

RELATIONSHIPS.- This species is readily separated 
from P. t oruiosus by its fifth sucker, b~y its somewnat smaller 
i nead ana smaller suckers. Tne ovaries of this species are more 
slender tnan in P. toruiosus. ?. l ongicollis is readily differentiat- 
ed from P. percae by its larger nead, larger s ackers, snorter cirrus- 
poucn, fewer testjiSjby its rewer uterine poucnes, and by the position 
or the genital pore. 


P. long-icon is resemoles P. cernua e in a v ry lew 
particulars sucn as short<©G/s.. of cirrus-poucn. However P. 
longicoilis nas a larger head, larger suckers, fewer uterine pouches 
smaller emDryos, lower ana smaller testes v/hich are differently 

P. longicoilis is much larger than P. fall ax and P. 
dub/C^/s , and it differs -further in having fewer uterine pouches, 
a shorter eirrug-pouch, fewer and larger testes, which are 
arranged in two fields. The genital pore in P. longicoilis is 
furtner anterior tnan in__P. fall ax or P. dubius . 

P. longicoilis diff ers iron P. Y)£q/etfus in having lev/er 
uterine pouches, fewer ana larger testes and a very aifferent ar- 
ragement 01 testes. 

P. longicoilis does not closely resembly any iorms yet 
reported from Horth America. 




(/V.3 ?r/*ya-^y 




Gmelin, 1790:3079, No. 79. 


Halysi s 


Zeder, 1803:376. 



ocellata ' 

Rudolphi, 1810: 108, in part . 




oc el lata 

Siebold, 1835:83. 




ocell ata 

Vcn Rat z, 1897:155,162. 




f ilicollis 

Vcn Rat 2, 1897:155,162. 




f ilicollis 

Vcn Rat 2, 1897:155,162. 

SPECIFIC DIAGNOSIS.- Characters of genus Strobila skort, 
robust. Observed length up to 40 pa. Max. "breadth 1.50 mm. Numb 
of proglottids 50-60. Segmentation not plain. Head not well set 
off from neck, flattened dcrsovent rally , and with a flat anterior 
face. Breadth of head 0.291-0.316 mm, length about 0.10 mm. 
Suckers not prominent, directed anteriorly, placed above broadest 
zone of head t Suckers almost globular, heavily muscled, small, 
with deep cavities. Diameter of suckers 0,064-0.090 mm, Eifth 
sucker present, 0.024 mm. in diameter, Neck 0.3-0.39 mm. broad, 
1.7-2.0 mm. long. 

First prcglottids much broader than long, 0.425 mm. 
broad x 0.085 mm. long. Mature and ripe prcglottids broader than 
long. Length greater than breadtn only in old ripe proglottids. 
Dimensions of mature proglottids about 0.476 mm. long x 0.816 mm. 
broad, ripe proglottid 1.309-1.51 mm. broad x 0.68-0.85 mm. long. 
End-prcglott id present and functional. 


GENITAL ORGANS,- Common genital sinus situated near middle 
of lateral margin of segment, irregularly alternating. 

Ifal e organs.- Cirrus-pouch. 0.185-0.228 mm. long, ex- 
tending barely thru vitellaria. Ratio of length of cirrus-pouch to 
"breadth of proglottid 2:9 in mature, 1:5, 1:6, 1:7 in ripe pro- 
glottids. Ductus ejaculat criu s in i 1-3 coils. Vas deferens 
forming a narrow mass extending to middle of proglottid. Testes 
about 70 in number, in one layer, occupying entire field between 
vitellaria and anterior to ovary. Liameter of testes 0.085-0.130 

F emal e organs . - Vagina anterior tc cirrus-pouch., never 
crossing the latter. Sphincter vaginae and receptaculura semimis 
not seen. Lobes of ovary thick, heavy and long, length of pair 
0.510-0.918 mm. Vitellaria coarsely follicular, voluminous, 
situated farther from r&argin of proglottid than in most species. 
Uterus in ripe proglottid s with 6, 8, 9, 12 lateral pouches on 
either side. 

EGGS.- Embryo with three-layered shell, second shell 
0.037-0.04 mm. in diameter, embryo 00212-0.0265. mm. in length. 


HABITAT.- In intestine of Acerina cernua (Linn) 





MATERIAL.- The description of this species is b a 3 GQ c n 
five specimens which Dr. H. E. Ward secured from Prof. Max. Brann. 
The material bears the label (Prot eocephalus ccellata Rud. int. 
Acerir cernua L. Kc'r igsberg , Pr. 8/93." 

These specimens bearing the number 09.26 and slides of the ! 
same are now to be found in Dr. H. B. Ward's collection. 

EXTERNAL CHARACTERS. - In general appearance these worms 
are short, robust. Four of them are very short, 10.0-19.5-22.0 mm 
The fifth measured about 40 mm. Yet each strobile has proglcttids 
with ripe eggs. A max. breadth of 1.50 mm. was observed in the 
longest specimen and a breadth of 1.309 mm. in a shorter one. 

The head (PI. 3 figJ.yj,y£ ) is net well set off from the 
neck. Indeed it is continuous with the neck which in all five 
specimens is broader than the head. The anterior face of the head 
is flattened. The suckers (PI. 3 figs.V^j^ ) are set at the 
margin of the flattened area.. They are small and not prominent and 
are directed anteriad. No furrows or ridges mark the head. A 
fifth sucker is very faintly visible. The head is 0.291-0.316 
mm. broad at a point a little posterior tc the suckers. It is 
flattened dcrsovent rally . Its length which cannot be determined 
with any accuracy t is about 0.100 mm. 

The suckers are heavily muscled, deeply concave, small 
i and nearly globular ir. shape. Six suckers, four from one head and 

two from a second , gave the following measurements: i 




0.07 mm 

0.079 mm. 
0.070 " 
0.064 ■ 
0.079 " 
0.085 " 
0.090 » 

Diameter of opening 

0.042 mm. approx. 
0.042 " 
0.037 " 

0.037 " 
0.037 " 

The range of length and breadth of the sucker is about 
0.064-0.090 mm. A sucker in transverse section measured 0.074 mm. 
in diameter. The sucker opening , it will be observed tis very small 
0.037-0.042 mm. 

The fifth sucker which seems to be a true sucker is about 
0.024 mm. in diameter. Its cavity is very shallow. Because of 
scarcity of material no sections were made and its true structure 
could not be finally determined. 

The neck in all five specimens was broad and thick. It 
is narrowest just posterior to the head. Here it measures about 
0.3-0.39 mm. Its length varies from 1.36 - 1.7 - 2.0 mm from the 
tip of the head to the first traces of segmentation. 


The total number of proglottids in one of the shorter 
strobilas was 52, in the longest, 66. The segmentation is not 
plain. The edges of the chain are quite smooth except for now and 
then small folds which rarel} r occur at the junction of two proglo- 
ttids. The angles between proglottids are scarcely noticeable. No 
longitudinal folds or furrows were seen. 

The first proglottids are much broader than long. In one 
specimen they measured 0.425 mm. broad by 0.085 mm. long. As the 
proglottids become cider they increase both in length and breadth. 
In all but a very few ripe proglottids the breadth exceeds the 
length or the segments are nearly quadrate. Mature proglottids are 
broader than long about 0.476 mm. long by 0.816 mm. broad. Pipe 
proglottids measure up to 1.309 mm. broad by 0.68 mm. long or even 
up to 1.51 mm. broad x 0.85 mm. long. One ripe proglottid which 
had discharged some of its eggs measured 1.19 mm. long by 0.915 mm. 
bread. A true end proglottid pointed at the posterior end, was 
1.19 mm. broad at the anterior end and 1.105 mm. long. It was 
functional. The measurements given are representat ive ; hence it 
will be noted that in general thebreadth of the proglottids ex- 
ceeds the length. Only rarely and then only in old ripe proglottid^ 
does the length exceed the breadth. The nervous, excretory 
systems, and the musculature have not been studied. 

GMITAL ORG/iTS.- Thecommon genital sinus (PI. 3 figs. A ) 
■ 3 situated very near the middle of the lateral margin of the segment. 
It may be slightly posterior to the middle in ripe proglottid. It 


alternates irregularly. There is no genital papilla. 

I* the worm of 52 proglottids the anlagsi of the sexual 

organs in one specimen of 52 proglottids appear in the 10th-12th 


segment as two darkly staining the one representing the 

cirrus and vagina and the other the ovaries. In the 20th segment 
the anlage* 1 of the testes appear. Beginning with the 34th segment 
the sexual organs are mature while with segment 40 a number of e^gs 
begin to appear in the uterus. 

Male organs .- The cirrus-pouch (PI. 3 figs.^yi ) in 
mature proglottids is about 0.221 mm. long while in ripe proglottids 
its length measures 0.185-0.212-0.228 mm. It is elongated ovoid 
in shape being broadest near the inner end. In mature proglottids 
it extends just a. short distance thru the vitellaria. In ripe 
proglottids it may barely reach thru the vitellaria. The ratio of 
the length of the cirrus-pouch to the breadth of the proglottid 
varies from 2:9 in mature to 1:5, 1:6, 1:7 in ripe proglottids. 

The prtruded cirrus has not been seen. Within the cirrus- 
pouch the distal portion of the cirrus is slender and straight. 
The ductus ejaculat orius is thrown into a few coilSjfrom 1-3. 

The vas deferens in proglottids scarcely mature is 
thrown into coils which extend in a straight narrow mass nearly to 
the middle of the proglottid, never past the middle. In ripe 
proglottids ceils of the vas deferens are massed at the inner end 
of the cirrus-pouch. There is no large mass of coils in the middle 
of the segment. In P. f allax the coils of the vas deferens form 
a dense mass in the midfield of the segment. In P. neglectus the 



large mass of coils of vas deferens lies in the middle of the 
proglott id . 

The testes (PI. J fig. H~} ) about 70 in number, ap- 
parently lie in one layer. They are irregularly scattered "between 
the vitellaria anterior tc the ovary. They measure 0.085-0.130 
mm. in diameter. They are thus more numerous and larger than in 
P. f 81 lax and P. dubiu s , and somewhat more numerous and larger 
than in P. percae and much larger than in P. neglectus . 

Female Organs .- The vagina (Pi. 9 figJ.y^y? ) always 
opens anterior to the cirrus-pouch which it never crosses in its 
course to the interovarial space. It passes to the middle of the 
proglottid in a smooth gentle curve, then it bends posteriad to- 
ward t^e inter- cvarial space. There are no ceils of vagina an- 
terior to the ovary but at that place it may be slightly sinuous. 
This straight or nearly straight condition of the vagina is evi- 
dence that the worm is in a normal state of contraction for 
usually when a proglottid is unduly contracted the vagina is very 
sinuous while in an elongated proglottid of the same worm the 
vagina may be straight or nearly so. 

Thus far in my material T have been unable to demonstrate 
a sphincter vaginae. The lumen of the vagina from its opening to 
the ovary is almost constant in diameter. JTo receptaculum seminis 
has been demonstrated. 

The ovary (PI. 3 fig. f7 ) is bilobed as in other 
JJroteocephalids. T^e lobes are thick, heavy and long. In all 


dimensions they are much larger than in P. f allax or P. dubius . 
They also have a different outline. In mature proglottids they 
measure 0.510 mm. long while in ripe proglottids their span may 
be as much as 0.918 mm. This is much greater than in P. f allax 
or P. dubius and more nearly like the condition in P. percae . 

The organs within the interovarial space have not been 

The vitellaria (PI. 3 fig. ) are coarsely follicular 

lateral masses. The follicles are much coarser and more numerous 
than in P. dubius or P. f allax . The vitellaria are situated at a 
distance of 0.100 mm. from the margin in mature and ripe proglottids 
This is considerably farther than in p, f allax and P. dubius and 
greater than in P. percae . 

The uterus a median tube in mature proglottids is 
augmented in ripe proglottids by 6 - 8 - 9 - 12 lateral out- 
pocketings on either side. r The pouches (PI. 3 fig*y2,y/) are 
long, reaching to the vitellaria, and are quite voluminous. Some- 
times the larger 'pouches are apparently subdivided by short septa. 
For this reason the appearance of the ripe proglottid in toto 
preparations differs considerably from that of P. percag or P. dubius 

EGGS.- The embryo is surrounded by three membranes, an 
outer thin and hyaline, a middle thick and somewhat granular, and 
an inner very thin membrane immediately investing the embryo. The 
embryo varies from ovaidal to sperical in shape. The measurements 
of four embryos may be taken as fairly representative of the range 


of size -- 0.0265 x 0.024 mm, 0.0212 x 0.0212 mm, 0.0265 x 0.0212 
mm, 0.024 x 0.0212 mm. The second membrane e measures 0.037-0.040 
BOB. in diameter. The outer membrane is very variable in size and 
in my material almost impossible to measure on account of collapse. 

RELATIONSHIPS.- This species somewhat resembling P. percae 
and P. t C30.QSUS in its outward appearance is distinguished from 
the latcer by its fifth sucker, its smaller suckers, its shorter 
length, its much mere numerous lateral uterine pouches, its fewer 
testes, and by the different arrangement of testes. 

Prom P. percae it is distinguished by its more numerous 
uterine pouches, its mere numerous and larger testes, its much 
shor er cirrus-pouch, and by the different relations of cirrus- 
pouch and vagina. 

Prom P. duhittg and P. f allax it i s distinguished by its 
larger bead, its proglottids of greatly different proportions, 
its more numerous and larger testes, by the greatly different 
cirrus and cirrus-pouch, by the different relations of the cirrus- 
pcuch and vagina, by the differently placed coils of vas deferens, 
by the differens in the size of ovaries, and by the shape of the 
ovarial space in the r:>pe proglottid. 

Prom P. longi colli s as described by Von Linstow P. cer^uae 
is distinguished by t"He much shorter neck, the different position of 
the genital opening, by the number of testes, by the position and 
shape of the mass of coils of the vas deferens ,by the number of 
uterine pouches, and by the size of the eggs. No American species 
yet discovered resembles it closely. 


TMs species is closely allied to the form which Zschokke 
desigratec as P. long; i col J. is (Pud) hut which I have determined to 
be a "ew species P. negj ectus . Unfortunately the heads of P. cernuae 
and _P. neglectus cannot be compared due to lack of heads in my 
Material of the latter. 

The shape and size of the proglottids may be about the 
same. The segmentation is evident in P. neglectus and very 
indistinct in this species. The minimum length of the cirrus- 
pouch of P. neglectu s is greater than the maximum of the same 
organ in P. cernuae n. sp. The ductus e jaculatcrius in P. neglectu 
is straight but it has from two to three ceils in this species. 

In ripe proglottids of P. neri ectus the ratio of the length of the 

to the proglottid breadUn" 
cirrus-pouchy is 1:3 to 2: 9 but in P. cernuae it is 1:5, 1:6, 1:7 . 

The coils of the vas deferens are massed close to the 

cirrus pouch in P. cernuae but they are mostly median in P. neglectu s 

The testes of P. c ernuae a re nearly double the size of those in P. 

negl ectus so also the length of the ovary in ripe proglottids is 

much greater in the former than in the latter. The extent and 

compactness of the vitellaria are different in the two species, 

likewise the relation of the vitellaria to the proglottid margin. 

There may be more uterine outpecket ings in P. cernua e, than in P. 

neglectus . The embryo measures 0.026-0.0265 mm. in P. n eglectus 

and about 0.021 mm in spherical embryos of P. cernuae . A maximum 

measurement of 0.0265 mm may occur in the latter species only in 

elongated embryos. These species which seem much alike at first 

glance are thus after all quite different. 



(PI. i Figs. ^3, fr/^J 

SPECIFIC DIAGNOSIS.- Characters of genus. Complete 
trobila, dead, suckers, nock not observed.. Outline of proglottid^ 
rougn. Segmentation evident. Genital pore on a slight prominence. 
Ripe proglottids U.o mm. long x mm. broad to u.7b mm. long 

x mm. broad. Young and mature p -oglottids not observed. 

GENITAL ORGANS.- Genital pores irregularly alternating^ 
situated near middle or lateral margin^cirrus and vagina opening 
into common genital atrium. 

Male organs .- Cirruspouch frequently contracted, O.ltfb 
-0.365 mm. long x U.UbU ram. broad. Normal cirrus-pouch U.340 mm. 
long, cirrus and ductus e jaculatorius in cirrus-pouch straight; 
vas deferens a mass of coils in midfield. Testes about 7b, spheroida 
u.u4ii-y.U64 mm. in diameter. Testes in one 1 aye r f filling space 
between vitellaria. 

Female organs .- Vagina anterior to cirrus-pouch. 
Vaginal opening dorsal to opening of cirrus-pouch. Sphincter vaginae 
poorly developed, situated near vaginal opening. Vagina crosses 
cirrus-poucn near middle. Receptacuiura seminis not observed. 

Ovary bilooed^postenor , looes with smooth outlines, club- 
snaped. Organs of interovariai space not observed. Vitellc.ria lateral 
follicular^ follicles small, compacted. 

Uterus with 7-y lateral pouches on either side. Uterine 
pores not seen. 


EGGS.- Embryo O,, 'db-<J .v^bb mm. in diameter, outer membrane 
U .042-0. 04 7 .mm. 

TYPE.- Bottle material oy.lo in Dr. .H. B. Sard's collection 
and slides from the same material. 

H A3 I TAT.- "Foreile" ( Tnutta rano Improbably irom Lake 
Geneva or Laice Lucerne, Sv/itzerland. 

MATERIAL.- Some pieces oi" strobila without heads v:ere 
received by Dr. H. B. Ward from Prol'essor Zsottokke at Basel. These 
v/ere labelled " Taenia longicoliis Rud. , aus Foreile, Zdohokke." 
The bottle is now Ho. u' in Dr. H. B. V/ard's collection. Four 
toto preparations uniiat"i.ened / v/ere made i'rom this material. The 
more slender parts or the strobila were not examined. 

EXTERNAL CHARACTERS.- Eead ; suckers , and neck were not seer 
A proglottid with eggs in the uterus measured 0.5 mm. long by u,93 
mm. broad. A ripe proglottid very lull oi' eggs was u.7b mm. long "by mm. broad. The outline of the proglottids was rough with severf 
a|l indentations on each proglottid margin. Tne limits of proglottids 
are well defined by the indentations at the corners and "by the 
rounded corners. A slight prominence bears the genital pore. 

GENITAL ORGAITS . - The genital pores are. irregularly 
alternating in the strobila, and situated one near the middle of 
the lateral margin eacn proglottid. This genital pore leads into a 
common atrium into which the cirrus-pouch and vagina open.. 

Male organs.- When the cirrus is partially protruded the 
fcirrus-pouchi is constricted near its inner end (Pi. *j fig. frj ). 


It is also reduced in length, measuring in various states of 
contraction form 0.185-0.190-0.220-0.265 ram. Its maximum breadth 
is about 0.60 mm. In a single proglottid the cirrus-pouchy was in 
a normal condition. Thf5 cirrus-pouch measures 0.390 mm. loner. 
The ratio of the length of the cirrus-pouch to the breadth of the 
proglottid is 1/4-1/3. 

The cirrus is sometimes protruded a very little. It is 
blunt at the tip. Within the cirrus-pouch the cirrus and ductus 
ejaculatorius are straight. The one passes imperceptibly into the 

The coils of the vas deferens cannot be clearly seen but 
their confused outlines may be discerned in the middle region of 
the proglottid left vacant of the testes. 

The testes, counted in two proglottides, numbered 75 and 
76. They are spheroidal, 0.042-0.053-0.064 mm. in diameter. They 
lie in one place apparently and fill the space between the vitell- 
aria, anterior to the ovary. 

Female organs . - Stained proglottids showed 7-9 large 
uterine outpocke tings on either side. So uterine pores were dis- 
covered. The vitellaria are I'ollicular and situated in the lateral 
fields of the proglottid quite near the margin. The follicles are 
small and are quite compacted. The vitellaria are much less heavily 
developed than in P. cj^v^ 

The ovary is bilobed and is posteriorly situated. The lobes 
are long, thick and heavy, and have a smooth outline. In ripe pro- 
glottids the span of the ovarian lobes may be 0.60 mm. 

The vagina is situated anterior to the cirrus-pouch but its 

opening into the common genital atrium is directly dorsal to the 
cirrus-pouch. Hear its opening it has a spincter vaginae 0.016 mm. 
lone? by 0.010 mm. thick, formally in its coarse. 


tie vagina is slightly bowed posteriad across the middle of the 
cirrus-poueft. In those cases in which the cirrus is partly protruded 
the vagina crosses tne inner end or the cirrus-pouch. The course 
of the vagina cannot to be traced to the interovarial space. The 
organs of zhe interovarial space have not been studied. 

EGGS.- The embrpo measured u.u2t>-u.U2bb mm. in diameter. 


The outer membrane measured U.U42-U.U47 mm. Tne second membrane was 

This species in some respects resembles J?# f allax 
which Zschokke had determined to be T« ocellata . The triangular 
shfipe oi" the interovarial space in ripe proglottids is very 
similar in both species. The proglottids are as a rule broader out 
shorter in this material. The uterine outpocketing are much alike. 
In the number of the testes and their close proximity to each other 
I this species is very different from P. fall ax .The cirrus-pouch is 
relatively shorter, but of much the same character. The vaginae of 
the two species in respect to sphincter are Buch alike. The ovary 
is quite sin"iar,tno considerably larger in this species. The testes 
and cirrus-pouch and the size of the embryo are good characters 
for differentiating the one from the other. 

In some ways this species resembles P. d ubuis but the 
ripe proglottids are a little larger and somewhat tnicker. The 
cirrus-pouch in proglottids of the same stage of development may 
be of nearly the same length. This species has mor«. testes wnich 
are smaller than in P. dubuis . The chief difference lies in the 
size of the embryo. This difference is from0.C0b-U.uu7 mm. The 


The second egg membrane in P. dub^rij s is larger than the 

outer membrane of P. n^ffleotu s» 

This species difiere from P. percae in the length of 
"Che cirrus-pouch proportion oi" the proglottids, number and size 
of testes, and in the size of the embryo. 

P. negiectus differs from P. longicollis in the number, 

The arrangement of testes . , ^ 
size, and arrangement of test s, in two fielas (?) in P. l onglcollis 

sets that species apart irom all other species reported from fish. 

It further difrers from that species in the number of uterine 

pouches and in the pos tion of the genital pore. 

P. negiectus i^oes not agree v/ali with Zschokke's 

(l8d4:l4-ib) description of Taenia longicollis Rud. from Coregonus 

f era. That species is apparently identical with the P. fallax 

from Coregonus f era 


PR0TE0CEPHA1US OSCU1ATUS Goeze) sp. ina. 
(Pl.<? figs. 
1782; Taenia o soul at a Goeze, 1732:415 

1SS2: Taenia alternate transverse l in^ata Goese, 1732:416. 

1786: Taenia o soul at a Batsch, 1786:209 

1786: Taenia siluri Batsch, 1786:157. 

1788: Taenia o soul at a Schrank, 1788:47 

1788 { Taenia nrlanis Schrank, 1788:47. 

1790: T aenia percae B. Gmelin, 1790:3079. 

1790: Taenia siluri Gmelin 1790:3080. 

1803: Halysis siluri Zeder, 1803, 353. 

1803 : Halysis Transve rse- lineata. Zeder 1803:353. 

1810 -. Taenia o sculata Rudolphi ; 1810, u. 2 ptz. :116 

19: Taenia o oc lata Rudolph! : 1819 ; 150 : 49 7 . 

1845: Taen ia osculata Dujardin, 1845:584. 

1850: Taenia osoulata Diesing, 1850:522-23 

1854: Taenia osculi.ta Wagener 1854. 

1896: Iohthyotaenia osculata Riggehbach. 1896:267. 

1909: Ichthyotaenia osculata Luhe, 1909:30-31. 

SPECIFIC DIAG1I0SIS.- See Luhe diagnosis at the end of th 

Host \ 





Jilurus r lanis 

He gdebfirg? : 


Goeze, 1732:415-16. 

Jilurus glanis ; 

Greii'swald : 


\ Rudolphi, 1819: 150 • 


HISTORICAL SUMLIARY.- Goeze (1782, P. 415.) described and 
figured this form from the intestine of Silurls glanis . His eiagEOSi 
is here given. 

Der gemundete 3andvnmn. Taenia ofculata. 
Auch too Graf von Borke in den GedaVmen eines 2 4p fundi gen 
We £es ( Silurus Glanis 1. ) ilach der GroJ^e dieses Fisches wenige, 
and auqjerst kleine Taenian. Kleiner, als in den Hechten. ITicht am 
llagenende , sondern mehr in der Gegend des Afters. Das Merkwiirdigst^e 
daran, das5 sie ausser den beaden gaugblascn am Kopfc , an dem t/orsteh 
endem Russel eine yvahrc Iluftdung hatten, welche unter dem K7*ompos*ito 
mit Ho. 4. Tub. 3. deutlich/zu sehen (var. An keinem andern hat der 
Graf die Russelmundiimg so deutlich, als an difen ivahrgenomraen. Sie 
waren alle lebendig* aber au^t der Glasplatte genw/er zu behandeln, 
weil sie sich bestandig kriimmten, und mit ihren Saugv/arsen selb^ 
an ihrem eigaem Korper vest ansogen. Ein besonderer Urns t and.' 1 
Das Aufblahen der §augwarzen, und das Austrdken und Einziehen das 
Russels ein angeneher^s Schauspiel. Dies Glieder zehv undeutlieh. 
Mehr Runzeln als Glieder. 

j Fi £* tJ Da - Xopfende des gorundeten Bandwurme aus dem 

leirt;, durcL Tub. B. L'o. 4. vergrosert* 

A. b. s'.vo oesonderc gcbildete Oa-agmundun-en: 
c d, die Oskula nit Stralen; 

e f , ein vorstehender Russel ; 
g, an demcelbenvorn eine Hudung: 
h i, der ungegliederte Hals. 
?ig. 10. Endrlieder , sehr re^^lrnasig -ezo ' <-.. ■.: -t i-r. c :lon c-. -nt an^xs. 




Eine wirklich neua and besondere Art. Auch aus den Ged- 


darmen eines seehzehnpfun&igen Wfelses { Silurus Giants). Als iff) to 
3sten Iulius 1780 von Rekane nach Magdeburg kan, Blieb ion daselbst 
urn der grosjeren Fische willen n^och einige Tage. I oh bekam die 
Gedarme (a^ejnes Stors (Aoipenser sturio I.) von; 50 >£)fund. In denselb 
versohiedene Askariden, und zv/een AWjer u/i a bea. andern Fisehen , 

Der MeohselswfrgjBe Linoirte Bandwurm. Taenia, alto mat Am 
transverse linoata: articulis qjiajgang alaribus ; oapite quadriverrusr 


In den Gedarmen eines fafelse^is ab er iand ion zween merkwurdige 

BanvTurmer • 


Die Jtange etd^ 12 Zoll. Die: Breite am breitestem 
ftinterende eine gute iinie. Die Glieder regelmasslge Quadrate: 
eine kinie aang und breit. Haoh dem Kopfe zu lTerhaltnismassig 
kleinev. Das Harakteristisohe dieges Wirms v/aren die, Weohselsweise , 
in jagem Gliede, in ungleioher Ordnumg ste^hende wejsse Cuj^erlinien, 
die bis uber ie Ha^fte des #6rpers, vom ^interevvde an, hinauf gehen, 
und besonders in die fcigen fallen. Diefe iinien gehen nicht weiter 

bis in dio ^itte des ^liedes. Zoigt sioh z. G. auf der @r/chten 

» « 
die linke Jeite trek* Dann auf der linken $eite in zv/es. Gliedern, 

und die reclite ,§eite freH. Weiter nur an einem Gliede, bald r coats, 

bable linkCs, u. s. u/. In der luittc. j e des (gliede s am jHlntcrende 

i'egon rti^e v£ier, unde ion kann diese Linien fur nichts anders, als 

fur Stergange halten, obgleich die Randmu'ndungen zu klein, oder zu 

verschloqjen S ind , als dayman sie Ivahrfe^hnen konnte. 


.Jfoj A'oflf^n&e sehr jfzin und dunnc , a"ber flaoh, und das ^6'pfch.en *\ 
opprtion wie oin kleines (fcotgen. Unter den fejfomposito der Hals 
ungegliedert . Am $ojjJe au# jeder < § e:3 '' te zwo stark hervorstehende 
JJaugroundun^gen.Vorn Gin fturae£ Russelchen, ohne Baken. 


Batsch, Sehrank, Hmelin, and Zeder described this species under 
various names (see quotation from Rudolphi 1308-1810 v. 2 pt. 2. p 
116) - 

Rudolphi (1808-1810 V. 2 pt. 2 p 116) gave a diagnosis, the 
synonmy, description and observations on this species. His descrip- 
tion is here given in full* 

Taenia osculata Goeze. 
Taenia : osculis rostellique apioe concavis . parte antica 
capillar i . articulis quadrat is planis , margine ma jorum integerrimo. 

Goese ITaturg. p. 415. Tab. 33. fig. 9. 10. T. osculata. 

Idem ib. p. 416. Tab. 33. fig» 1% • 14» taenia altematim transverse 


Batsch Bandw. p. 209. n. 45. fig. 146. 147. T. osculata. p. 157. n. 
16. fig. 80. 82. T. siluri. 

Sehrank Verse ichn. p. 47. n. 141. T. osculata. n. 142. T. glanis. 
Qmel. Syst. Nat. p. 3080. n. 82. T. siluri. ib. p. 3079. n. 77. T. 
percae p. 

Tabl. Encycl. t. 49. fig. 4. 5. (ic. Goez.) T. siluri, fig. 6-9. 
(.ic. Goez.) T. alternans. 

Zeder ITaturg. p. 353. n. 40. Halysis siluri. n. 41. Hal. transverse 

Hab. in Siluri Glanidis parte intestini praesertim media, saepe tamen 
etiam inter prioren. Goezius Iulio, ego Majo, reperimus. 

Deecr. Vermes pauci a Comite de Borke reperti exigui, duo 
a Goezio observati duodecim pollices longi; mei, copiosissimi, 
pollicem ad pedem et quod excurrit longi, ant ice capillares, postice 


lineam ad sesquilineam lati. 

Caput discretion, subglobosum, exig^uum, Oscula . uti caput 
totum, quam in specie praecedente multo minora, concava, profunda, 
ostio exiguo, ut in ilia sita saepeque simul in conspectuin venientia. 
Rostellum intermedium, brevissimum, osculum quintum, sed minus et 
plerumque magis protrusum refert. Corporis plani et tenuis pars 
antica in verme tarn pollicari quam pedali capillaris, mox tantum 
rugosa vel incisa videtur colli speeiem sistens, mox autern articulis 
distinctis iisque tenuissimis, forma variis, monilif ormibus, oblongis 
et rugae f ormibus inter jectis , constat; pars reliqua sensim incors- 
cens, articulis tandem quadratis, satis aequalibus margins lateral! 
rectiusculo, intererrimo. Articulos ultimus rotundatus, fere semper 
emarginatus. Cujusvis articuli majoris media pars ovarium, sive 
maculam oblongam, pellucidam continet, quae neque anteriorem, neque 
posteriorem articuli marginem at'. ingit. A quolibet ovario linea 
transversa ad alterum articuli marginem lateral em, ordine plerumque 
altero, non tamen certo, excurrit* Linea longitudinalis , qualis in 
specie praecedente, in hac Hon concpicua. 

Substantia vermis tenuis, at firma et duriuscula, ideoque 
in aqua non pro speciei praecedentis more intumescit vel gelatinosa 
fit. Vermibus utuiusque speciei aeque magnis, prioris volumen multo 
ma jus . 

Obs. 1. Hujus et praecedentis diBcrimen, quo minus con- 
jungantur, nimis magnum esse, cuique observation en illi adjectam 
lwgenti, vel utramque recentem aut spiritu vini servatum comparanti 
patebit . 

Obs, 2. Me speeiem a Comite de Borke detectam et Goexio 


osculatam dictam, cum altera hujus auctoris specie (transversim 
lineata) conjunxisse fcrsan mireris, sed utrique eollum non articu- 
latum et tenue adscribitur, quale praecedenti semper deest, et B 
Borkius , qui Taeniolas suas in Glanidis intestinis crassis repererit 
nonnisi partem vermium summam offendisse videtur, dum articulos rugaj 
potius referre asserit, quod de hujus tantum parte antica valet. 
Goezius vermes majores reperit et equsdem ecor (fig, 11.). Taeniam 
integram sistens, bona est. Capitis verc ieones neque Barkiana 
neque Goeziana laude dignae sunt. Zederus, verme licet non viso, 
alteram Goezii speciem rescindendam esse, recte praedixerat. 

Obs. 5. Millie rus de Taenia ocellata, nobis n. 21. dicta, 
loquendo, Taeniam transverse - lineatam Goezii ab ilia lineolarum 
lateralium defectu solo forsan differre, suspicatus est, quo 
Gmelinus commotus, qui Taerdam nostram oeellatae varietatem fingeret 
sed osculata tcto ceelo diversa, et rostello solo jam distingui 
potest . 

Obs. 4. Omnes quidem Taeniae osculatae sunt, nostra autem 
(uti et praecedens) ita vocari potest, cum os- 

culo quinto quasi instruatur, et oscula valde proiunda sint. 
Goezii ergo nomen conservavi, et praecedentem simili voce disignavi. 

Rudolphi (1808-10 V. 2. pt. 2:115) also described Taenia 

calycina from Silurus glanis . This species he later (Rudolphi 1819 

497. ) came to consider as identical 77ith Taenia osculata. His 

observations are here given:- "Entozoologiam edando duplicem h&jus 

Taeniae formam, mihi turn temporis constantem visam prae oculis 

habui, quo commotus. T. calycinam mihi dictam ab osculata distinxi. 


Po::tmo::um vero Gryhiae Cctobri mense Siluri Glanidis quinquap-inta 


tres Hibras pondere aequantis intestina examinando, Taenias 
plurimas reperi inter T. osculatam et calycinam et ita quidem in- 
termedias, ut nullum amplius discrimen superesset. Tres quatuorve 
pedes lon.Tae collo mox longiore mox breviore utebantur; articuli 
lineis longitudinalibus , rarius ( posteriores ) etiam transversis 
insignes; aquae commissae quasi gelatino sae reddebantur, aliquot 
dies vivae conservatae . Taenia calycina ideoque rescindenda est." 

Dujardin (1845) and Liesing (1850) added very little to the 
previous diagnos£s anc descriptions. 

Carus (1867) gave tv;o drav/ings of this form. Riggenbach 
(1896) considered it a species of Ichthyotaenia "but he gave no 
descriptive data on the species. 


hUhe (1909:30-31) gave a short descriptive diagnosis of the 
species and in an original figure, reproduced (PI. f Fig. /fo ) 
he shov/ed a little of the structure of the proglottid. His diag- 
nosis reads: 

r/ Zirka 50 cm "bis 1 m lang, bei einer grossten Breite von ca. 
3 mm. Scolex mit einem kleinen scheitelstandigen "Saugnapf", der 
mit zahlreichen, sehr hinfalligen Hakchen von 0,007 mm Lange 
besetzt ist. Proglottid en mit entwichelten Geschlechtsdrusen sov/ie 
auch solche mit Uterus annabernd quadratisch. Genitalof f nung in 
der ;.:itte des Gliedrandes. Cirruabentel klein, kaum uber den 
Dotterstock nach innen hinausragend ; das Inauel das Yas deferens 
stark quergestreckt , vcm Cirrusbentel bis an die Lied: anlinie reichencl 
Hodenblaschen sehr zahlreich, das genze I.:ittelbeld der Proglottis 
einnehmend. Vagina ohne Sphincter. Jteruy jederseits mit etwo 
6-8 schlanken, sich et?;as verzv/eigenden Scitenasten. "Calkkorperchen 


arscheinend sparlich. 

Im Barm von Slluras glanis L." 
SSnce the papers by t* Linstow- (1891J-.anC. Llonticelli (1891) 
this species has been closely associated "by mariy with that proup of 
cestoces of fish which has been variously known as Tetracotylus 
Monticelli, Ichthyotaeni a Lonnberg, and Proteocephalus We inland # 
Until the present those generic names have been considered to be 
synonyms, Then it is proposed to limit the genus Proteocephalus it 
is difficult to determine the systematic position of some of the 
less well known species among which may be mentioned Taenia osculata 

I recognize that this species differs from typical species 
of Proteocephalus in the possession of spines in the fifth sucker 
and perhaps for that reason it should be placed in another genus or s 
new genus should be erected for it. But I do not consider that 
sufficient accurate data are available to justify such action. 1 9 
therefore, retain it in the genus Proteocephalus where it must be^^' 
|»s a species inquirenda. A careful study of good material might show 
that this species belongs in the genus Tetracotylus llonticelli. 

i (n 



t 21* 5 figs./^-J/3j 

. SPECIFIC DIAGNOSIS.- Characters of genus. Strobile short 
and slender. Observed length up to 90 mm. Max. "breadth 1.24 mm. 
He ok 3-7 mm. long "by 0.20-0.25 mm. broad. Proglottids, first in 
chain very short, 0.05 mm. long by 0.250 mm. broad. Mature and ripe 
proglottids nearly quadrate or in a fev; ripe proglottids length 
exceeding the breadth, (J>. 595x0. 595 mm. up to 0.730 mm. long x0.595 
mm. broad. End-proglottid present, functional. Segmentation not 
very distinct, intersegmental furrows shallow. 

Head conical, flattened dorsoventually, without furrows, 
showing great variations in shape. Breadth of head about 0.33, 
thickness 0.22 mm, length 0.20-0.25 mm. Suckers cup-shaped, deep, 
muscular, 0.095-0.105 mm. in diameter. Aperture of same 0.06-0.07 
mm. Fifth sucker present, muscular, with shallow cavity. Diameter 
of fifth sucker 0.05-0.075 mm. 

GENITAL O RGANS .- Common genital sinus irregularly 
alternating at or near middle of lateral margin of proglottid. 

MALE' ORGANS.- Tests ovoidal, 0.05 mm. long by 0.04-0.05 
mm. broad, 54-70 in number in a single layer taking up field 
between vitellaria anterior to ovary. Ducutus e jaculatorius nearly 
straight. Cirrus well muscled, straight, when protruded 0.07-0.08 
mm. long, 0.02-0.025 mm. broad. Cirrus-pouch short, stout, 0.13- 
0.14 mm. long by 0.05-0.06 mm. broad, length into proglottid breadth 


FSMAT.y. -.''^SAIS t Vagina usually anterior, but vaginal 
| opening always dorsal to cirrus-pouch. Vagina crossing inmer 
end of cirrus- sheathe, phincter vaginae small. Receptaculum 
i small, situated anterior to ovary* . 1.4kus when fully developed 
with 10-14 lateral pouchtj on either side. Uterine pores on 
ventral surface, 2-3 in number. 

EBGS . - Embryos 0.016-0.018 mm. in diameter. 

HABITAT: In intestine of 


: Locality 

: Collector 

: Authority. 

Esox Reticulatue 
Le 3eur : 
Esox Lucius Liifi 

i Sebago Lake , Hair 
■ W Inut Lake, Hi cl* 

le H. B. Hard. 
i.T.L. Hankinson 

: La Rue . 
: La Rue. 

Esox Lucius L^-nit .uake Geneva, 7/is< A. J. Coats. * La Rue. 

TYPE.- Alcoholics in Dr. H. B. Ward's Sebago Lake Col- 
lection llos. 4,6,7, 3, 9, 107. Slides of same. Material collected 
from Esox Reticulatus Le Seur (Type host), Sebago Lake, Me. 
(type locality) 

COTYPE Alcoholics Ho. 413 in T. H. Hankinson'a 
collection, from Jsox Lucius Linn, Walnut Lake, Michigan.. Slides 
of same. 


AUTOTYPE . - Alcoholics ITo. 10.35 in collection Dr. 
H. B. 7/ard from Esox / acius Linn, Lake Geneva, Wisconsin. 

HISTORICAL SUMMARY.- Leidy (1888: 169 and 1SJ0: 417) 
described some specimens of cestodes from Esox Reticulatus. The 
two specific names which he proposed for them T consider to he 
synonyms and have included under the one name . Proteooephalus 
ftematosoTrtus (Leidy). Loidys specimens were considerably larger 

than specimens of P. ^>inguis and apparently are not the same 
Fo<- a more complete discussion see page 


SOURCE OF MATERIAL.- This study is based, for the 
most part, upon specimens collected by Dr. H. B. Ward from the 
intestines of Esox ^3ticulatus Le Seur at Sebago Lake, Maine while 
engaged in an investigation of Sebago Lake for the U. S. Bureau of 
Fisheries, summer of 1907. Portion of the study are also based upon 
s ec 'mens secured by Prof. T. L. Hankinson from Escx 7^ucuis Linn, 
in the course of a biological investigation of Walnut Lake, Llich. 
during the summer of 1906. This last material was referred to by 
La Rue (1910: 25) as " A new species, which I shall describe in 

detail later " It bears the number 413 in Mr. Hankinson V 


. Dr. H. B. Y/ard examined six specimens of Esox Reticulatus 
Le Seur while at Sebago Lake. These six specimens yielded sixteen 
specimens and pieces as follows: No 4, 1 head 2 pieces; !To. 6, 2 
heads 2-3 pieces; No. 7-1 head 2 pieces; No. 8, 11 heads 2 
pieces; No. 9, 1 head; No. 107, 5 heads, 1 young cestode, 10 
large pieces. In many instances the specimens reported as heads 
were complete worms while in others the end proglottid and part 
of the strobila were missing. The-inf ect: on of Sebago Lake was 

Hankinson* s material in bottle No. 413 from Esox tucius 
of V/alnut Lake was prcbable from a single host. It contained 
60-70 specimens of P. ^ingu&s . 


A "bottle Ho, 10.35 in Dr. H. B. Ward's collection, 
labelled "from stomach and intestine of a"Pickerel" Lake Geneva, 
Wisconsin, Jy. 1, 1904. A J. Coats, collector " contained 5-6 
complete spec mens of P. p inane s and several pieces of the same 
pecies. Prof. George 7/agner of the University of V/iscontin states 
repaying to a letter of inquiry that only one species of pickerel, 
the Bsox fcicuis linn; is found at Lake Geneva, Wisconsin. 

The data on this species were studied from specimens 
cleared and examined in glycerine, from specimens stained and 
mounted in toto and from transverse an >. frontal actions. 

EXTER1TAI CHARACTERS.- These cestodes are small 
and slender r ( pi . 5 f ig./o£ ) , The largest specimen from the Sebago 
Lake material measured 90 mm. long "by 1,24 mm. in max. breadth. 
The longest specimen from the Walnut Lake host measured 60 mm. 
long by 1.0 mm. in max. Breadth. The strobila of a 50 mm. 
specimen is made up of an unsegmented neck 3-7 mm. long by 0.20- 
0.25 mm. broad followed by about 300 proglottids. The youngest 
proglottids measure about 0.050 mm. long by 0.250 mm. broad. 
Further along in the chain these increase in length and breadth 
until in mature and ripe proglottids the length and breadth are 
about equal. Measurements of several ripe proglottids are as 
follows, the length in mm. being stated first: 0.595x0.595, 0680 
xO.630, 0595x0.663, 0.730x0.595 mm. T$e proglottids are broadly 
r llipsoidal in trans-section. 


A typical end proglottid^with a rounded posterior 
end is present. The end proglottid is functional. The segmentation 
is not very plain. The intersegmental furrows are not deep. The 
corners of the proglottids do not project. The lateral margins are 
somewhat rounded and smooth. 

1^,101,101', I'lilM 

The head (P1.5" figs^/s ) is a cone shaped structure somewhat 
flattened dorsoventr ally, without folds or furrows. Its "breadth is 
about 0.33 mm. Its thickness about . .22 mm. its length about 0.20- 
0.25 mm. In extreme cases the head may attain a breath of 0.45 mm. 
This excessive breadth is due to the contraction of the longitudinal 
muscles of the head and neck. A collection of these specimens shows 
heads of very different ctatesof contract i or . The head bears on 
its broadest par"'" four Leep cup-shaped muscular suckers, wQaich 
are directed forward and outward. On its apex the scolex bears a 
well formed, muscular, fifth sucker. 

The diameter of the sucker ranges from 0.09 5-0.105 mm. 
thile the diameter of the aperture varies from 0.06-0.07 mm. The 
fifth sucker measure 0.05-0.075 mm. in diameter. Tho its cavity 
is quite shallow it is a function sucker as is shown by its 
typical musculature (PI. f f ig/<*) . 

INTERNAL AHAT0IJY.- The cuticula ana musculature have 
not been found to differ in their essential features from those of 
other members of the genus. The nervous system in the head is 
#ttch like that oi p. Filaroides described by me. A ganglionic 
mass (PI. f fig. /&$-) is situated at the level of suckers. Two 
main lateral nerve trunks extend thru-out the length of the strobila 


Ho accessory nerve trunks were observed. 

Excr e Sory System .- The excretory system in the head is 
made up four ;ain longitudinal canals which at various levels 
receive many anastomosing coiled vesLels. These pass thru the 
tissues of the scolex and neck in every direction. In the strobila 
j there are two pains of lateral excretory vessels, ventral and 
dorsal respectively. These lie within the medullary parenchyma. 
The dorsal vessel passes dorsad, the ventral vessel ventrad to the 
cirrus-pouch and vagina. In diameter of lumen the ventral vessel 
greatly exceeds the dorsal. The membrane lining the ventral 
vessel is thin while that lining the dorsal ves. el is thick. 

A posterior excretory commissure in each proglottid 
xjould not be found. Numerous secondary excretory openings which are 
connected with the ventral vessel and less frequently with the 
dorsal vessel have been seen. In the end-proglottid the four main 
excretory ducts jdischarge to the exterior by a common pore. A 
bladder could not be s en. 

GEHITA1 OROAJNS.- The common genital sinus is marginal 
situated at or near the middle of the proglottid. While the vagina 
frequently lies anterior to the cir -us-pouch, the opening of the 
i vagina without exception is dorsal to the opening of the cirrus. 
There is no genital papilla. 

MALE ORGAIS.- The test's f Pl.jT f IgJoi ) are numerous, 
54-70. These are in a single layer in the medullary parenchyma 
occupying the entire field between the vitellaria anterior to the 
ovary. The teste* measure up to 0.050 mm. long by 0.04-0.050 mm. bro»d 

============ — 


The vasa efferentia (PI. 5 flgjw) form a network of fine 
anastomosing tubules just dorsal to the testes and within the 
medullary parenchyma. The main branches of the vasa efferentia 
unite to form the larger vasa deferens The junction occurs at a 
point near the median line about one-fourth the distance from the 
anterior to the posterior and of the segment. 

The vas deferens from this point of juncture forms a 
mass of coils extending to the cirrus-pouch. This mass of coils 
functions as a vesicula seminalis. 

When the vas deferens enters the cirrus-pouch it rto®tv©§ 
some additional glandular elements the prostate cells. This part 
of the vas deferens is called the ductus e jaculatorius. The 
ductus is very nearly straight. 

The cirrus-pouch is elongated oval i$ outline. It is 
short and stout being 0.13-0.14 mm. long by 0.05-0.06 mm. broad. 
The ratio of its length to the proglottid breadth is 1:4 to 1:3. 

Female Organ r . - The vaginal pore opens into the 
genrtal sinus just dorsal to the cirrus-pouch tho the vagina it- 
self may be anterior to the cirrus-pouch ; (/f/'*~ //o, 
A small sphincter vaginae is situated near "Sie openfng^m iifield 
the vagina crosses the inner end of the cirrus-sheathe. V/hen 
passing posteriad to the intefovarial space the vagina is straight 
not sinuous nor coiled. Just anterior to the ovary the vagina 
dilates slightly to form a receptaculum seminis. 

The organs of the interovarisO. space are typical of the 

genus . 



The ovary is hilobed. The lobes ( Pl.jf f ig./oi ) are 
elongated oval, smooth in outline. The vitellaria are typical 
of the genus. 

The uterus is formed of a ventral-lying tube which extends 

nearly the full length of the proglottid. From this tube by the 

process described by me for Pj_ ^ilaroides lateral outpo eke tings 

arise. These lateral pouches ( PI. f figJ^) number 10-14 on either 

side. As these lateral pouches beco^v: distended with eggs the 

outlines of some of them are nearly obliterated. Yet in all but the 

ripest proglottids the full number is visible. Sections thru 

mature and ripening proglottids show the pouches in all stages of 

development. Sections also show the ventral diverticula which 

finally come to open on the ventral surface. Such diverticula. 

were described by me ( 1910:36) for P. ffilaroides and their 

presence in other species was at that time noted. I n P. / »ingu t s 

the number of ventral diverticula is greater than Gui Schneider 
P. m a r, c r o c o p 1 1 a ljg , j? . percao . ?. esocis . 
• (1905! reported i n^P . Ambigaa but smaller than the number which 

occurs in P. yilar rifles . A drawing (PI. f fig./flf) of a frontal 
section just below the cuticula show two of these openings. In 
such sections the openings, usually two or three, and the tubes 
leading up to them could be readily traced. A specimen cleared in 
glycerine showed these openings beautifully. The actual count 
of openings on 25 consecutive proglottids is as follows: 1, 2, 3, 
2, 2, 2, 2, 2, 2, 3, 2, 2, 2, 2, 8, 3, 2, 4, 3, 2, 4, 2, 3, 3, 3,. ' 

EGGS.- The uterine eggs were observed only in 
pre served condition. The outer mucilaginous envelope could not 
be ^een. 


The embryos measured 0.016-0.018 mm. 

RELATIONSHIPS.- P. ^ o inguis is much larger than P. 
e socis ( Gui Schneider). It has a fifth sucker which the latter 
lacks. It iiffers from that species in having a larger head, 
larger suckers, larger proglottids more tests, and a relatively- 
shorter cirrus-pouch. 

P. p inguis differs from P. ^usillus (Ward) which occurs 
in the same region in having a larger head, larger suckers, longer 
proglottids which are of different proportions from those of P. 

usillus . The ,egmentation is of different character. The 
relative lenjfch of the cirrus-pouch is very diffewnt. 

P. Singula iiffers very much from P. exiguus in size, 
in the proportions of the proglottids, in the size of the head and 
of the suckers, in the relative length of the cirrus-i. ouch, in the 
character of t"ie uterine pouches and in the size of the embryos. 

P. finguis differs from P. n<acrocephalus in having a 
fifth sucker, in being considerably mailer, in the relative length 
of the cirrusgouches in the number of testes, and in the size of the 

P. pinguis is smaller than P. gmbloolitis , P. a ingu/af is 

and P. ferplexus . In the character and size of the hei i and suckers 

there ij also considerable difference. It is further differentiated 
from those species by the presence of a fifth sucker ./Mch those 
species lack. 


P. j gingols is quite different from those European 
forms which have*fifth sucker. These species are P. . .;rcae . 
P. Sailor, P. ^uhius , P> tsmuae . P. /on-ic. .llis * It differs 
from them in size of "body of suckers, and proglottids, in relative 
length 01 cirrus-pouch in the size of the embryo and in the 
number of tests . 

1 u 


(PldFigs. J**f3^ 

SPECIFIC DIAGNOSIS.- Characters of the genus. Cestcaes 
imall, length 30-50 mm. Max. "breadth 0.350 mm. Proglottid? few. 
Segmentation distinct. Head spheroidal, frequently much contracted, 
breadth about 0.300 mm. length about C.260 mm. Four suckers placed 
at broadest zore of head. Suckers, deep, 0.140 mm. long x 0.110 mm. 
broad. Fifth sucker well developed, muscular, 0.060 mm. in diameter. 

Heck 1-1.5 mm. long by 0.21 mm. bread. First proglcttids 
broader than long., breadth of same 0.090 mm. Mature prcglottids 
longer than broad. Pipe proglcttids much longer than bread. Length 
of sane 0.84-1.4 mm, breadth 0.18-0.35 mm. End-proglott id present 
and fertile. 

Genital organs .- Genital si^us marginal, irregularly 
alternating, situated at end first 1/3-2/5 of proglottid. Vagina 
usually anterior and dorsal to cirrus-pouch, rarely posterior. 

Male organs .- Testes 44-60-70 in number arranged in two 
layers between vitellaria. Length of sane 0.06-0.096 mm. breadth 
0.04-0,05 mm. Vas deferens a small mass of coils anterior to cirrus- 
pouch. Cirrus-pouch, 0.095-0.106 mm. long by 0.053-0.060 mm. broad, 
reaching l/3-l/4 across the proglottid breadth. Ductus ejaculator- 
ius forming one or two coils in c irrus-pouch. Girrus short and 
straight . 


Female organs . - Vaglnia never crossing cirrus-pouch. 
Sphincter vaginae and Receptaculum seminis not seen. 

Ovary bilobed, posterior, Lobes shcrt, thick free ends of 
same pressed posteriad. and together. Vitellaria sparse, follicles 
small . 

Uturus with 10-14-16 lateral pouches on either side. 

EGGS.- not observed. 

ILABIT/.T . - Intestine and oesOophagus of Sftlmo sebfrgo Girard 
(typshost) SehCigc Lake, Lie. (type locality) and intestine and 
pyloric c'&ecx^ of C rist ovomer nam ay cush Walbaum, Lake Temagami, 
Ontario . 

TYPE.- Bottles 15, 16, 42 in the collection from SebUgc 
Lake, Me, summer 1907, and slides fro-a this material. 



The following description is quoted from Ward (1910:1185- 

87) . 

P:'ct eocephalu s pusillus nov.spec- Adult cestode with 
short, strobila, measuring only 30 to 50 mm. in length. Proglottids 
scanty, segmentation distinct. Head, much contracted. Neck 1 to 1.5 
mm. long by 0.21 mm. broad. First prcglottids 0.09 mm. broad, 
changing gradually until in mature proglottids the length greatly 
exceeds the breadth. Hipe proglottids measure 0.84 to 1.4 mm. long 
b- 0.18 to 0.35 mm. broad. Terminal proglottids present and fertile 
Sexual organs typical for Prot eocephalus ; uterus meclian, with 10 to 
14 lateral out pocket ings on either side. Testes numerous, within 
vitellaria. Genital pore lateral, one-third to two-fifths of length 
or proglottid from anterior margin of same. Ovaries bilo ed, median 
isthmus indistinct, anteropost ericr diameter nearly equal to breadth 
of both lobes. Only a few specimens obtained from a single host 
species, Sal mo sebago . 

This species approaches most nearly to P. ocellata and P. 
percae among known species. Unlike the new species, however, both 
of these cider forms have a fifth sucker, fewer lateral uterine out- 
pocketings, a longer neck, differently shaped ovaries, and markedly 
different proglottids. 

In specimens with developed proglottids the head was so 
much contracted or distorted that any special description would be 
of little value. One could easily observe the general features 
characteristic of the genus. There was no well developed terminal 
or fifth sucker, and the end organ, which is known to replace it in 


■any forms of this genus, was inconspicuously developed, if present. 
Personally, I incline to the view that on more careful examination 
this structure will be found in all species, even these in which 
its absence has been made a matter of record. Accordingly, not 
much weight can be put in its presence or absence in any individual?' 

Three plerocercoid larvae or ycung cestodes were found in 
company with P rot ecce phalus pusillus, which I regard as young forms 
of this species. The largest came from the salmon which was most 
heavily infected with this cestode parasite. It was 3.15 mm. long 
and had begun to assume clearly the appearance of an immature 
cestode. The head measured 0.3 mm. wide by 0.26 mm. long, and the 
suckers 0.14 mm. in length by 0.11 mm. in width. The neck was 
slightly narrower than the head, but was not clearly set off from 
the body, which was very uniform in diameter and measured 0.25 mm. 
in average width. The posterior end of the body was swollen into 
a rounded knob about 0.35 mm. broad and of approximately the same 
length. This feature was evidently produced by a powerful contrac- 
tion of the terminal region of the body. In and near it one could 
see very indistinct indications of proglottid formation. In form, 
size, and general aspect this young cestode was in full agreement 
with the anterior rep-ions of the mature cestodes of this species 
with which it was associated. The head, which, was not contracted, 
shewed on careful study the delicate outline of a rudimentary end 

or a mere definite discussion of this peculiar structure so 
variable in development in the cestodes of this genus, I would refer 
to a paper now in press by my student, Mr. George R. LaRue, to whom 
I am indebted for. a comparison of this material from Salmc seb g,gp 
with preparations of other species of Prot eccephalus . 


organ. While such a structure was not demonstrated in the mature 
individuals described above, one can say positively that if present 
it could not have been seen owing to the greatly contracted condition 
of the adult scolices. I believe that its presence will be demon- 
strated in more favorable specimens. The complete agreement of this 
largest larva with the mature specimens in all other features compels 
me to regard both as different stages in the development of the 
same species. 

The other larvae were still in early stages of development 
and probably had been ingested by the salmon at a very recent date. 
Their relationship is not so clear in all respects, and yet I do not 
hesitate to associate with the new species of Prot eocephalus a 
plerocercoid or young cestode obtained from the same host as the 
adult worms and the older larva just described. The head is broadly 
conical, without furrows, and measures 0.3 mm. in breadth. The 
suckers measure 60 to 74/* in diameter. There is no rostellum or 
fifth sucker to be found, while the end organ is so poorly developed 
a to be visible with difficulty and only under the most favorable 
circumstances. The reck is nearly as broad as the head. In general 
appearance this larva resembles the adult cestode and the cider 
larva previously described. With some reserve one may also assign 
to this species a single plerocercus taken from another specimen of 
Salm o s ebago . The head, which measures only 150/" in breadth, is 

shaped like that of the young cestode and like it is without rostell- 
um or fifth sucker, while the end organ is difficult to demonstrate, 
"either furrows nor ridges are seen on the larva, which has a. total 


length of 1.14 mm. The sucker measures only 30 to 45/x in diameter. 
The neck is slightly narrower than the head. This form certainly 
belongs to the genus Proteoc ep halus and probably to the species 


already described. 

Dr. H. B. Ward has very kindly loaned me his specimens 
for study. Additional preparations have been made and from them I 
have been able to secure additional data on this species. 

The type material, bottles 15, 16 and 42 was taken from 
the intestine and oesophagus of Salmo sebago Girard, Sebago Lake, Me 
Jy. 27 and Aug. 6, 1907. Other specimens cf the same species were 
taken by Dr. H. E. Ward from two specimens of Cristovcmer j^amaycush 
Walbaum, Lake Temagami, Ontario in August 1911. These last are 
described separately. 

EXTERNAL CHARACTERS.- This is one of the smaller species 
of Prot eocephalus . The observed length is from 30-50 mm. for 
specimens which have ripe proglottids. The proglottids are few in 
number and the segmentation is distinct. 

The head is spheroidal. In many cases the hesd is greatly 
contract ed, so much so that its structure can not be determined. 
The head measures 0.300 mm. broad by about 0.260 mm. long. The four 
suckers measure 0.140 mm. long by 0.110 mm broad. The sucker cavity 
is quite deep. At the apex cf the head (Pl./fig^) is a fa(r$ly well 
developed fifth sucker which in well stained specimens shows an 
evident musculature and a cavit.v. Ward (1910) reported only an 
end-organ instead of a fifth sucker. When his specimen was restained 
and mounted it shewed a very characteristic fifth sucker. The fifth 


sucker measures 0.060 mm. 

The neck is 1-1.5 mm. long by about 0.21 mm. broad. It 
passes over quickly into the first proglottids which are broader 
than long. Their breadth is about 0.090 mm. The proglottids rapid- 
ly become quorate or even longer than broad. Pipe proglottids 
measure 0.84-1.4 mm. long by 0.18-0.35 mm. broad. An end-prcglottid 
is usual] y present and fertile. 

GENITAL • ORGAMS . - The common genital sinus is marginal, 
situated at the end of the first 1/3-2/5 of the proglottid. The 
position of the genital sinus alternates irregularly in the strobila. 

Female organ s .- The vagina (PI. (fig!*) usually opens 
tntericr and dorsal to the cirrus-pouch but sometimes (P]/.fig'.p it is 
posterior to the same. The numbers of proglottids examined have been 
too few to establish whether the posterior position of the vagina 
is a normal occurence. Even when in the anterior situation the 
vagina does not cross the cirrus-pcuch. From its opening it makes 
a long curve tovvtrd the middle of the proglottid which it reaches 
at h point posterior to the inner end of tJs.» cirrus-pouch. The 
vagina is dorsal to the uterus. P~ vagina.1 sphincter could not be 
demoBstrated. If present at all it must be very .eak and perhaps 
quite long. I have not demonstrated a receptaculum seminis. 

The bilobed ovary (Pl/>f igjty'yis peculiar in that the lobes 
which are short and very thick are pressed closely together. This 
is a condition not yet found regularly in any other species of the 
genus. In transverse sections the thickness and compactness of the 
ovary is noted at once. The vitellaria are sparse. The follicles 


are small. 

The ducts of the interovarial space have net been thoroly 
investigated. An obtype and an occapt are present. 

The uterus (PI ./,f ig'.^) in ripe proglottids shows from 10- 
14-16 lateral outpocket ings on either side. The uterine pores have 
not been seen. 

Male o rgans .- The testes (PL|fig.i*) number from 44-60-70. 
Since these lie in two layers the figure does not show all of them. 
In less mature proglottids where the testes are much smaller and 
more distinct they can "be counted more readily. The testes measure 
0.06-0.072-0.096 mm. long "by 0.040-0.050 mm. broad. 

The vas deferens (PI ./ f ig. ,<J ) fonns a small mass of coils 
in the middle of the proglottid. The larger part of the mass is 
anterior to the cirrus-pouch. This is a condition not usually met 
with in this genus. 

The cirrus-pouch which is rather heavy reaches 1/4-1/3 
across the proglottid breadth. Its length is 0.095-0.106 mm. and 
its breadth 0.053-0.060 mm. 

The ductus ejaculatori us forms a coil or two within the 
cirrus-pouch before passing over into the short straight cirrus. I 
have not seen the cirrus protruded. 

On account of a lack of sufficient material but very few 
sections been made and these were too poor to yield much 
valuable data. 

The eggs have not been observed. 

Other specimens which I have provisionally assigned to thi 


species were taken by Dr. F. B. Ward from the pyloric and intestinal 
rerions of Cris t ivomer yiamfly cush Walbaum, Lake Temagami, Ontario, 
August 1910. These specimens "bear the lot numbers T>P, T<j,Ti/j, vli P, 
in Dr. H. B. Ward's collection. 

EXTERNAL CHARACTERS .- The specimens were small measuring 
up to 20-30 mm. long. Wo specimens had ripe proglott ids- When the 
worm is fully mature it is perhaps considerably longer. 

Measurements of a number of specimens yielded the follow- 
ing data: Average breadth of 7heads 0.284 mm., Max. breadth of 
s&me 0.340, minimum 0.244; average length of 6 heads 0.177 mm. max- 
imum length of sane 0.238, minimum 0.133. Average length of 17 
suckers 0.134 mm. maximum length of same 0,179 mm., -777/711 mum 0.106,* 
average breadth 0^ 13 suckers 0.108, maximum breadth of same 0.127, 
minimum, 0.096 mm; Sucker opening 0.058-0.080 mm.; fifth sucker 
present; diameter about C.060 mm. Average length of 6 necks 1.49 mm 
maximum length 1.70 mm. minimum, 1.02 mm; Keck narrow, up to 0.135 

The proglottids are few, 34-40. First prcglottids are 
broader than long, or quadrate, length soon exceeds breadth, and 
mature proglottids are longer than broad measuring up to 0.56-0.68 mm 
long x 0.24-0.265 mm. broad. The maximum breadth of 0.340 mm was 
observed in a proglottid measuring 0.357 mm. long x 0.340 mm. broad. 
Segmentation is very distinct. 

GENITAL ORGANS.- The genital pore^is near the middle, 
usually anterior thereto but sometimes posterior. The vagina opens 


anterior to the cirrus-pouch. 

The lobes cf the ovary are thick and bent back until they 
nearly meet in the median line. The vitellaria are sparse. The 
fully developed uterus has not been observed. 

The testes are in two layers. They are irregularly 
scattered between the vitellaria and anterior to the ovary. The 
testes number from 44-66. They measure 0.159x0.180 mm. In many 
proglottids the testes are not quite as fully developed as in the 
specimens from Salmo S ehago so they are more readily counted. 

The mass cf ceils of the vas deferens is small and it 
lies largely anterior to the cirrus-pouch. The ductus ejaculations 
forms a single coil within the cirrus-pouch. The cirrus is not 

The cirrus-pouch extends to the middle of the proglottid or 
nearly thereto. The length of the cirrus-pouch varies from 0.111- 
0.159 mm. with an average of about 0.133 mm. for eight pouches. The 
breadth ofthe pouch is abcut 0.053-0.070 mm. Figs, were drawn from 
these specimens which came from Crist ^ivomer nar ra ycush . 

The most serious discrepancies between the two lots is 
in the length of the cirrus-pouch and in the size of the suckers. 
The discrepancy in the case of cirrus-pouch is a matter of but a 
few micra and it may be caused by different states of contraction 
or by age. 

The suckers of the material from Cristivome r nar; a /cush are 
no larger on the average than those of the typical P. pusillus but 
in one or two cases the maximum length and breadth of the sucker is 


quite a little larger. It is to "be remembered that the measurements 
of the suckers of the typical P. pusillus are based on a single 
head. ' Consequently the range of size in that species has not been 
det^mined. Tho no ripe proglottids were present in the material 
from Cri st ivomer nam aycush and consequently some of the best 
diagnostic features could not be determined, it seems best to con- 
sider these forms identical. 

Pro teocephalu s pusil lus . Ward is closely allied to P. 

exiguus but is differentiated from that species by its larger head, 
larger suckers and larger sucker openings. The measurements of 
proglottids of the two species are much alike. However, the type of 
segmentation is very dissimilar. In the reproductive organs P_j_ 
pusillus varies from P. exiguus in having more and larger testes. 
The testes of P. pusillu s lie in two layers while in P. exiguus they 
are in one layer. The cirrus-pouch of P. exiguus is :uch longer 
than that of P. pusillus. The ovaries of the two species are very 
different. In the one species the lobes are slightly arched while 
in the other the lobes are bent nearly together. The lateral pcuches 
of P. exiguus are larger and Scarcely as numerous as in P pusillus. 

P. pusillu s somewhat resembles P. f alloX but is smaller. 
The head and suckers of P. pusillus are larger than those of P_. 
f allox . P pusillus has more and larger testes, a much shorter 
cirrus-pouch, and more uterine outpocket ings. The ovaries of the 
two species are very different. 

P. pusillu s differs from P. agonis B arbieri in having a 
fifth sucker. Moreover P. pusillus is a larger form than P. agonis. 


P. agon is has smaller and more numerous testes than P. P^i llu _£i an(i 
also fewer uterine outpock et ings . 

P. pusillu s differs radically in size and proportion from 
P. percae (0. P. Miiller) . The differences between the proportions 
of the proglottids and the size and relations of the genital organs 
are very marked. 

P. pusillus differs from P. esocis (Gut. Schneider) in 
having a larger head, larger suckers, in possessing a fifth sucker, & 

a shorter neck. The proportions of the segments are different. 

and the 

The length of the cirrus-pouch ratio of its length to the proglottid 
breadth are very different in the two species. The character of the 
segmentat ion is very unlike. 

There is very little probability that P. pusillu s is the 
same as P. sa l v el in f Linton. P. Salvelini has no fifth sucker. Its 
head and suckers are larger than those of P.. pusillus . The cirrus- 
pouch of P. salvelin i is also larger than that of P. pusillus . 
Linton's description of P. salvelin i does not furnish many character 
for determination and his drawing of the proglottid is so indistinct 
that it cannct be relied upon as a source of data. 


( PI. / Figs, l-f ) 

16^5: Baonia ma rorocephala 

1845; Taenia macro cephala 

1350: Taenia macrocephala 

1860: Taenia macrocephala 

1875: Taenia macrocephala 

1886: Taenia dilatata 

1 93: Taenia macrocephala 

SrepUn, 1825: 
Dujardin, 1845:585. 
Diesing, 1850:513-14, 
Molin, 1860:13. 
V. LinstoiV, 137, . 
Linton, 1886:438-9. 
Olsson, 1893« 

oh a /a 

l ohthyotaenia macro cga Rigfrenbach 1896:267, 

1897: Taenia dilatata. 

189 7: Taenia ifocrocephala 

1893: Taenia macrocephala 

1898: Taenia macrocephala 

1901: Taenia |)ilatato 

Linton. 1897:425. 
Stossieh. 1897:7. 
Stossieh, 1898:113-14. 
Muhl ing . 1893:373-69. 
Linton 1901: 435. 

1903: lohthyotaenia hemisphagttea G. Schneider 1903:29. 

1905: lohthyotaenia macrocephala G^ Schneider 1905:10,17-1!) 

1905: Proteocephalus macrocephalus lufer 1905:134-152. 

1909: lohthyotaenia macrocephala Luhe, 1909:33-4. 

SPECIFIC DIAGNOSIS.- The characters of the genus. Adult 
cestodes with strobila up to 40 cms. long, breadth 1-1.8 or perhaps 
more. Segmentation quite distinct$. Proglottids very numerous: first 
much broader than long; mature, broader than long or nearly quadrate; 
ripe, longer than broad. Neck 50-70 mm. long, 0.1-0.25 mm. broad. 
Head , globose, flattened, dorsovenut .^lly, 0.28-0.30-0.32 mm. broad 


0.15-0.16 ram thick. Suckers 0,095-0.106 mm in diameter. Fifth 
sucker replaced by end organ 0.025 mm in diameter. Sex organs as in 
genus. Genital aperture near middle of proglottid, vagina opening 
anterior and dorsal to cirrus-pouch. Testes 100-120 irregularly 
scattered between vitellaria lying in 1-2 layers usually. Uirrus- 
pouch short, about 0.160 mm long. Ratio of length of cirrus-pouch 
to proglottid breadth 1/6-1/8. Uterine pouches 7-11-12-14 on either 
side. Embryos 0.017-0.019-0.021 mm. 


HABITAT: In intestine 

or Anguilla 

vu/ -jar is and A. 


Host : 

; Local itv 

i • 
i « 

> i»o±±ecoor « 

au tnor ixy • 

Anguilla vugiar 

-is Greiiswaid 

Crepiin : 

Crepiin (1825:by-7i 

it n 

: Rennes 


Dujardin ( 1845,: 585) 

Tt tt 


lioiin ; 

Moiin (ibbu:i») 

it it ; 

V. Linsoow. : 

Lins gov/ ( id 75 : J 

ft tr 

Oiisson (18y3i J 

M It 



Stossich ( 18y7:7 ) 

It If 



Stossich (18y8:ll3-i< 

tt Tt : 

Memel,E. Prussia Mining 

Lluhiing ( 18y8 : by-37 , 

it it * 

; Finland 

: Schneider 

Scnneider (iyu3:2 v J) 

: Lake Lucerne ') Nuier « 

» K 

Nufer ( iyU5: 134-152) 


^. Wood's Hole, Mass. 

: Linton 

: Linton ( 188b: 488-9 


: Sebago Lake, Me. 

: H. B. ".Yard. 

HISTORICAL SDMHARY.- Crepiin (1825) found this species 
in the intestine of Angoilla Vulgaris at Greifswald. His diagnosis 
is here given: 


^Taenia macrocephala:- T. Capite elongato, antioe crassiore, 

c seulis -globosis anticis, rostello breuiss, no. obtuso, collo brevi , 

articulis anterioribus breuibus abtasis, sequentibus subquadratis , 

I oram le mniseis margin alibus usage alternis. 

Hab. In int e s Anguillae Martio, April ma jo at 

»unio hanc Taeniam riperi ant singuluar out minimi tamen eopia. 

Descr. Vermes a laquot l'neas usque ad octo polices longi, 

najores medio corpore lineam e m guarta uel dimidic. ejus parte lati, 

jubdiaphani , albissimi 

Caput, oblongum, antiee latias, rostello brevisimo. 
obtuso instruct u m, inerme, et quatuor__osc_ulj : s ( quoru rn. oril'icia 
non uid:) globosis, prominentibus , anticis, instructam. Collum 
brevtes, rugosum, cum capite contimuu, A^ti^jyj, antici breuissirai, 
inacquales, obtusi, oossim medio constrict i, eeguentes sensira 
latitudine et longitudine crescunt, maximan partem guidem latiores, 
quara longi mament, demum uero quadrati et ultirai adeo panlo longior s , 

| quara lati euadunt. Ommee per articulos decarrit ad margines lateral 
utrinque l inea alba^opaca. I orarnina articulorum Posticorum 
marginalia Ufe-^e a Herna, ex/quibus $ir-rhas progendet br-epfis, 
subtilise imus. 

Obs.i. Repori prirao hujus uerm is specimens uhicam valde 
parvum d. 27 mart. 1822., postea d. 23. Apr. ej. anni herara oarua 
specimina una cum Bothriocephalis elavaecipi tibas it idem paruis 
domun uero soeciemen octopollicare d. 23, majil822 et secundum 
specimen maynum d. 5 Tunii 1823 simu 1 om. illi adjacente. Bother 
clauaecipite edioeri V: uos examinar e microccopii opo mon mihi 


bicuit, at ma jorem deseription is pertem deverme in spireita dare debuerim. 

Obs. 2. Specimen June 3 intyentum uivaeissimum, ajgue- 
friftidae immissum, valdopere se movebit et contor^uebat , hisque 

otibus maxumum ovorara copiam demittebst subglobosorum, at tot 
a a^ua inde tarnida i'ieret. 

Obs. 3. Hujus Vermis caput sine d*ibio 0. F. Multer (Vid. 
Schriften 3er. 1. Ilaturf , Ii p. 208. cit. in Rudolphi. Entosoologiae 
I. II P. . p. 29. ) vidit, cum Bothriocephalo clatfaecipiti( Taeniae 
--.uguillae sibi dicto, a nostra taenia omnibus notis diversissimo ) 
capttt osculis quatuor instructum adscriberet . ; ' 

Dujardin (1845:585) described and reported specimens of 
this species thus: " — Long de 8 a 220 mm, large de (?) a 3 mm, 37;- 
tete allongee, plus epaisse en avent, avec une trompe tres-coiirte , 
obtuse; — ventouses globuleuses, dirigees en avant; — cou court; — 
premiers articles courts, obtus, les suivants presque carres, avec 
lea orifices genitaux irregulierement alternes, et les penis courts, 
tres-minces pendants.'' 

II. Creplin l'a trouve plusieurs fois, a Greifsv/ald, dans 
L'intestin de L'anguille. 

— J'ai trouve aussi a Kennes, dans une anguilie, trois 
jennes tenias longs de 6 mm et 10 mm et 13 mm, 5, larges de mm, 25 
a mm, 3, ayant la tete large de ram, 33 a mm, 48, sans trompe, 
et les ventouses larges de mm 106, dirigees en avant. 


Diesing (1850:513-14} added nothing to the earlier des- 
cription. IJolin (1860) reported a specimen of this species from 
Anguilla vulgaris taken at Padua in March 1859. Von Linstow (1875) 
described very insufficiently P. macrocephalus from Anguilla vulgaris , 
His diagnosis reads: "Descr. Vermes aliquot lineas usque ad octo 
pollices longi, majores medio corpore lineam cum quarta vel dimidia 
ejus parte lati, subdiaphani, aTbiscimi. 

Caput oblongum, antice latius, rostello brevissimo, obtuso 
instructum, inerme, et ouatuor osculis (quorum orificia non vidi), 
globossis, prominent ibus , anticis, instructum. Collum breve, rugosum 
cum capite continuum. Articuli antici brevissimi, inaequales, 
obtusi, pasim medio constrioti, sequentes sensim latitudine et long- 
itudine cresunt, mazimam partem quidem latiores, quan longi manent, 
demum vero quadrati et ultimi adeo paulo longi ores, quam lati evadunt 
Omnes per articulos decurrit ad margines laterals utrinque linea 
alba opaca. Foramina articulorum posticorum marginalia vage alterna, 
ex quibus cirrhus propendet brevis, subtilissimus. " 

Von Linstow 's measurements of the suckers much exceed 
those given by G. Schneider and myself for tlis species. He also re- 
ported smali fifth sucker that was very difficult to find. I am in- \ 
clined to believe that von Linstow actually saw the small end organ 
which in some Proteocephalids can rarely be seen in toto preparations, 


Linton (1886:438-9) described a speoies from Anguilla 
ohrysypa Raf., caught on our eastern coast as a new species, Taenia 
dilatata in these words: "Head small, truncate, or, in living speci 
mens, slightly prominent in front. Acetabula nearly circular, direc 
ed a little forward. Heck rugose, very long, very contractile and 
dilatable, narrow in front, tapering toward the head; a short dis- 
tance back of the head expanding into a number of irregular, trans- 
parent, dilated folds, which border both sides of an opaque central 
portion, in which two longitudinal canals are faintly outlined. 
First segments about three times as broad as long; median segments 
square, or broader than long; ultimate segments nearly square, some- 
times broader than long, sometimes longer than broad. Genital 
apertures marginal, opening a very little in front of the middle* 

A single specimen of this species of Taenia was obtained 
from the intestine of the Common Eel ( Anguilla vulgaris ) August 26, 
1885. The length of the specimen, when stretched out by fastening 
one end with a needle to the bottom of the dissecting dish and re- 
moving all kinds and curves with a fine brush, was 170 mm. The 
length of the same specinen, after having been preserved in aDcohol, 
is less than 90 mm. The specimen when first obtained and placed in 
sea-water was quite active. The body was constantly throwing itself 
into sinuous curves, while the head and neck were jerked from side t 
side with a moderately rapid motion. In addition to these movements 
the neck and anterior portions of the body constantly changed their 
shape by the inflation or dilatation of the investing membrances 
into wide transparent folds, constricted at irregular intervals by 
narrow transverse bands. The neck, meanwhile, was alternately 


stretched out and contracted like the body of a Nemertean. The 
anterior end of the head protruded into a proboscis like papilla. 
She breadth of the head itself varied from 0.17 mm to 0.35 mm. 

In the alcoholic specimen the dilatable folds of the neck 
are much contracted and broken. THe lie in rough, ragged frills 

along each side of the dark central part of the strobile. The 
head i r truncate of blunt in front. The neck immediately behind 
the sucking-disks is al ost as wide as the head, flate, thin, and 
little, if at all, tapering. 

The following measurements were made on the living specimen. 
The head and neck changed their position and shape so rapidly that 
it was with the greatest difficulty that trustworthy measurements 
could be made : 

Holl, Mass., August 26, 1885; one specimen. 

Von linstow (Compend. der Helminth., 1878) records but two 
T aenia e from the Common Eel, T. macrocephala Creplin and T. 
Eeraispherica Molin. T. di- -tat a is very different from the former. 


Breadth of head 

Diameter of acetabula 

Diameter of neck, narrowest part 

Distance of first segments from head 

2 ength of fourth segment from end of strobile 

Breadth of same, posterior end 

Breadth of same, anterior end 

Length of posterior segment 

Breadth of same, posterior end 

Breadth of same, anterior end 


Habitat.- Common Eel (Anguilla vulgaris); intestine; Wood's 


Liesing (Revis. der Oeph. , AB. Cycl. , p. 378) mentions the latter, 
"but gives no enumeration of characters. I do net have access to 
Uolin's caper, and cannot, therefore say whether di 1 at at a is 
identical with his species or not. The -eculiar inflated character 
of the neck suggests T. ambigua Dujardin, but the difference in 
size betv;een the adult specimens is alone sufficient to render 
their union in the same species impossible. 

Again Linton (1397:425) reported this species from A. 
chrysypa thus: Ho. 4812, J. 3. II. M. From common eel (Anguillq 
chrysopa). Several strobiles but in bad state of j reservation; 
no scolices; anterior ends have been exceedingly long and slender. 
The characteristic dilatations of this species can not be proved 
from these specimens. The general appearance of the strobiles, 
as well as the character of the segments, however, agrees with 
this species. 

It may be added that the segments of these specimens 

agree with Molin's description of his T. heraispherioa . With the 

evidence at hand, however, I do net feel justified in uniting the 

two species of T. dilatata and T_. hemispheri ca. 

Prof. H. B. 7/ard secured this material, no. 4812, If. S. 

IT. ~. for my examination. Its state of preservation was too poor 

to permit any positive determination being made. 

Linton (1901:435) recording the parasites of Anguilla 


chysypa reported this species again* "Taenia dilatata -iinton. - 


Specimens of this genus also taken in 1: 99 three in August two. 
Dimentions in millimeters: length, 8, diameter of head, 0.20 
diameter of sucker, 0.08 segments not mature. One specimen August 
20; length 14 mm. 3 

The folds on the neck are of no real diagnostic value. 
Sucj folds may he seen on may other specimens of cestodes in 
certain conditions of expansion and contraction. Unfortunately 
Linton's drawing of the mture proglottid is suc& that it permits 
no positive determination. His drawings of the head and of the 
form of the last proglottids are of value for comparative purpose. 
He makes more of those observations on the internal anatomy of 
the worm which are of great importance in the taxonomy of the group. 

-rinton "believed that the segments of his species agreed 
with Molin's description of. T, hemisphaerica . 


Kiggenbach (1896 J considered P. dilatatus (Linton) as identical 
with P. rlemisphaericus (Molin). This may he true if P hemi sphaer i cus 
(IJolinJ and P. macrocephalus (Creplin) are identical. ITufer (1905) 
thought P. dilatatus . P. hemisphaer icus , and P. macrocephalus iden- 
tical. Since I have had no material of P. hemisphaer icus 1 have pre- 
ferred to consider the last two forms as separate species* I am, 
however, ready to state that P. dilatatus is a synonym for P. 
macrocephalus . That name should therefore he dropped from the list 
of Proteocephalus species. 

Prof. H. B. Ward very kindly secured for my study Linton's 
slides and some alcoholic material of his P. dilatatus . These 
specimens from the statements in his letter to Professor 7/ard , must 
be from the lots described by Linton in 1886 and in 1901. Measure- 
ments of the heads of the suckers of the proglottids, of the cirrus- 
pouch, as well as of the general appearance of the v/orms, give me 
ample reason for considering this form to be identical with specimens 
taken from Anguilla Chrysyga . Lake Sebago, He, which I am describing 
as P. macrocephalus . 

Olsson (1893) reported the specimes from Anguilla vulgaris 
in Scandinavia. Riggenbach (1896) considered the form to be a 
species of I chthyo taenia . Stossich (1897:7) reported this species 
from Anguilla vulgaris at ITarenta, and again (1898:143-4) he gave 
a short diagnosis of specimens taken from the same host species at 
Trieste. In this diagnosis nothing of the inner anatomy of the worm 
is given. His diagnosis reads: "Lunghezza 220 mm; larghezza 3.5 mm. 
Scolice in continuazione del collo, allungato, ingrossato al- 
l'innanzi, con grandi ventose globose, situate anteriormente ; ro- 


stello cortissimo, ottuso, provveduto di una piccola ventosa apicale. 
Collo corto. Proglottidi prime corte e ottuse, le seguenti sub- 
quadrate. Aperture genitali irregolarmente alterne; pene corto, 
sottilis..,imo , pendente, con una strozzatura verso l'apice. Uova 
con due invogli, lunghe 0.089 mm. Rara nell* intestino dell' 
Ang uilla vulgaris (Trieste). 

Stossich was evidently describing P. macrocephalus and not 
the P. hemisphaericus fMolin). 

Mflhling (1898:37,69) noted this parasite in A. vulgaris in 
East Prussia. He gave no description. G. Schneider (1903:29) 
reported from an eel a single specimen of what he then identified 
as I chthyo taenia hemisphaeria . Later in a footnote (1905:10) he 
says of these specimens " Ichthyotaenia henisphaerica koramt, wie es 
scheint, in den aalen des Finnischen Meerbuse'ns garnicht vor. Das 
examplar welches ich fruher eimmal intumlich als hemisphaerica 
bestimmt habe (Beitrage zur kenntnis der Helminthenf auma des 
finnischen Lleorbusens, Acta soc. pro Fauna et Flora Fennica 26, no. 
3. 1903. S. 29), erwies sich bei genauerer Untersuchung auch als 
Ichthyotaenia macrocephala . 

From this statement I infer that G. Schneider at that time 
considered P. hemisphaericus to be a distinct species, 
j In the text of the same article Schneider (1905-17-19) 

described P. macrocephalus recording for the first time in the 
history of the species a careful study of its inner anatomy. For 

this reason a condensed resume of his description is here given. 


This species is clearly a close relative of P. £ercae 
and is differentiated therefrom by the very short cirrus-pouch and 
the short cirrus. 

The scolex has a diameter of about 0.300 mm. The diameter 
of the large deep suckers is 9.100 mm. The end organ is as in P. 
oercae a rudimentary structure made up of long cells and having a 
diameter of 0.025 mm. I have shown however that this organ in P. 
p_ercae is a true sucker. 

The neck is quite broad and straight compressed dorsoven- 
tvally. The length of the "body reaches 40 cms. and more. A 
specimen 11 cm. long and 18 mm. broad had about 200 segments. The 
youngest proglottids are about five times as broad as long. The 


middle mature proglottids are broader than long, ripe proglottids 
| art. longer than broad. 

The ovaries appear from the surface as two small stripes 
as in P. p_ercae.In old ripe proglottids they are of irregular shape, 
bent^ and somewhat overlapped. 

The cirrus and vagina open irregularly alternating right 
or left in the middle or a little anterior to the middle of the 
proglottid margin. 

The cirrus-sheathe is scarcely 1/6 as long as the proglottid 
breadth, often much shorter. 

Tne vas deferens ^cirrus-^ouofel forms some convolutions in 
the proximal end of the cirrus sheathe and here widens out into a 



tfesicula seminalis, in diareter 0.030 ram. The convolutions of the 
vas deferens without the cirrus-sheathe are eccentric on the side 
where the cirrus opens. In other species with a long cirrus-sheathe 
thls^convoluti on^ lies in the middle of the proglottid. 

The vagina has a smaller sphincter close to the opening. 
I* passes to the ovarii region without broadening out to a 


reeeptacuUlm seminis. The oocapt is very muscular. The uterine 
passage is 0.0;. mm. in diameter. 

The vitelline reservoir is 0.30 mm. in diameter. The 
vitellaria are voluminous, follicular. The opening of the uterine 
passage and the well preformed uterus opening are found almost in 
the same transverse section posterior the middle of the proglottid. 

The uterus has about eight branches in either siiffe. 
The testes which measure about 0.150 x 0.075 mm. lie irregularly 
in one or two layers. There are about 100 testes in each proglottid. 

Ventral excretory vessels are about 0.009 mm. in diameter, 
the dorsal vessels about CU)0^5 mm. This species occurs only in 
Anguilla Jolgari^s and always only in small numbers. 

Grepjiin's specimens were taken at Grei^swald in Prussia 
while Schneiders' were taken in Finland. Geographically these 
habitats are not distant. On this account as well as by reason of 
anatomical similarities it is extremely probable that Schneider's 
specimens were of the same species as grephinis. It is to be noted 
that Dujardia'jj observation on the size of specimens taken from 
Anguilla Vulgaris at Re\inos, France agree very well with those of 


Nufer (1905) described and figured what he considered to 
he P. macrocephalus Greplin. He found two specimens in the intestinal 
tract of a single ^mjoilla vulgaris from Lake Lucerne. Nufer shows 
such stricking inconsistencies in his statements which moreover 
agree so poorly with G. Schneider's and with my own, that it seems 
best to analyze parts of his paper. 

Compare his description of the head (p. 137) and his 
statement about the head in his summary(p. 145) with the facts which 
he presents in tabulated form (pp. 146-147) concerning P.macrocephalui 
P. £cell -tus, P A longicellis, and_P . toru\>asus . all of which species 
he reported from Lake Lucerne. 

"The Scolex is sehr gross, ziemlich breit , rundlich, und 
besitzt einen Durchraesser von 0*1512 mm. Gegen den hals ist er 
scharf and deutlich ab^esetzt und uber trifft alle ubrigen irn 
^ierwaldstattersee vorkommenjden Proteoce ohalenskolices an Grouse 

edeUtendes ( the i-^? lies are mine) Wesh^lb Creplin, 
auch auf den ^edftnken gekommen sein mag, diesen Proteocephalen mit 
Macro cephalus " zu bezeichnen, Vorne ist der Skolex mit vier grossen, 
yunde^n Sangnapfen versehen, die ihm die wuchtige Gestalt verleihen, 
wie ich sie sonst bei keinem Proteocephalen mehr angetroffen habe. 
Je noch der Kontraktion der Sagnapfe erscheint ihr Rand kreisrund 
Oder unregelra'assig gelappt. Der aussere j0urchnesser derselben 
betragt 0.069 mm, der innero Durchmesser 0.0414 mm. Da der §kolex 
bei meinem Praparate etwas geschrumpft ist, war es San ujpoglich; 
einen funften, scheitekstandigen Sattgnapf nochzui^eisen, wie ihn 
vnn T.i Tiatnw heohoichtet haben_willj .jedoch ist sicher. 


dass eine Scheitelveetiefung vorhanden 1st". 

Also in his conclusions note this (p. 145) , "Der skalex 
ubertrifft die ubrigen Proteocephalenskolices an Grosse; dement- 
aprechend ist auch der Durohmesser der Sangnapfe grosser". 

In the tabulated data (pp. 146-147) Hufer gives the 
dimensions- of the scoleces and of the suckers of the four species, 
thus; P. macrocephalus breadth of scolex 0.51 mm. diameter of 
suckers, 0.069 mm; P. ocellatus breadth of head 0.114 mm. 
diameter of suckers 0.05 mm. 0. longicollis , breadth of head 0.43 
mm. diameter of suckers 0.1E-C .19 mm; P. t orulosus , breadth of head 
0.255 mm. diameter of suckers 0,1 mm. 

According to these figures neither the diameter of the 
suckers of P. macrocepha lus not the breadth of the head exceeds 
those of any of the other species, except those of__P. ocellatus. 

Again note that in the last sentance of the first para- 
graph quoted llufer states that he could not find a fifth sucker, 
yet in his tabulated data (P. 146) he gives the diameter of the 
apical sucker as 0.026 mm. In this he must be quoting from von 
-instow's description. Evidently ITufer's statements lack accuracy. 

IJufer's drawings of the scolex are of littel help in 
giving a clear idea of the position of his specimens. 

Consider now his description of the male reproductive 
organs (pp. 140-142) and carefully compare his description with 
my figures (pi. 2 fig 35) 



Der cirrusbeutel ist keulenf ormig; vorne an der i.lundung 
schmal, erigreitert er sich nach hinten zienlich rasch unci erhalt 
ein&n grossten Durchmesser von 0,0945 mm. Am Hinterende ist er 
abgerundet und gent bis uber die Uitte des Gliedes hinaus. 'Vie 
von Linstov; (644 am Cirrus, so beobachtete ioh am aussern Drittel des 
Cirrusbeutels eine Einschnurung, durch die er ungefahr die v..n v. 
Linstcv angegebene gestalt eines Kegels erhal^ wie sie beim Kegel- 
spiel ublich ist. Er dringt nicht vollkommen gerade in das Innere 
des Gliedes ein, sondern richtet sich etwas schrag gegen des 
Hinterende der Proglottis. Die 7/andung des Cirrusbeutels wird von 
einer kraftig ausgebildenten. ..luskulc.tur umgeben. Aehnlicli v/ie 
bei Proteoce phal us o cellatus heft en sich an die Basis des Cirrus- 
beutels kraftih entwickelte itetrak^oren , ?;elc,e wie buschelige 
'Vurzelf asern an einem Bulbus sitzen. Der Raum zwischen der '.Van 
dung des Cirrusbeutels und dem Cirrus ist von zarten Bindegewebszelle; 
undzahlreichen, unregelmassig verteilten Parenchyrrkernen erfullt ; 
vielleicht sind auch einzelne kleine Zalkkorperchen eingestreut, 
wie sie von Xraer^er (55) bei Proteocephalus ocellatus gefunden 
worden sind. 

Der Cirrus bildet die direkte Fortsetzung des Yas de- 
ferens; er durchbricht den Cirrusbeutel am Hinterende und durchzieht 
inn in geradliniger. Yerlaufe. ilachdem er den hintern Drittel des 
Beutels passiert j'at, nimmt dessen Durchmesser zier.ilich rasch 
ab und reduziert sich auf die Halite. STaob den Beobachtungen 


Kraomers (53) ist der Cirrus be! Proteocephalus o ceilatus and P. 
torulosus in seinem jintern Teile in mehrere Schlingen gelegt, 

wahrend or bei P. macri'ceohalus als vollstandig gestrecktes Organ 
auftritt. Gleichbedeutenu v/ie diese ochiingen gelegt , Cirrus, 
dazu dient, "bei der Begattung den Cirrus zu verlangern. Bei dem 
Geschlechtsakte vrurde aann die ringf ormigen TJulste sich glatten 
und successive verschwinden in den Masse, als der Cirrus vor- 
gestossen v:ird. Er ist von einer strukturlosen, glatten wandung 
umgeben, der nioht wie "bei P. ocellatus und P. torulosus nach 
hinten gekrummte Chitinhakchen aufsitzen. Sein Lumen betragt im 
hjntern Drittel 0,009 mm, nimmt aber nach vorne bedeutend ab. 

Das vas deferens liegt hint or und vor dem cirrusbeutel 
etwas gegen den sitenrand des Gliedes verlagert und bildet ein 
dichtBn Knauel, der sich besonders in der Lange der Proglottis 
ausdehnt ; die Schlingen, die nit samenfaden erfullt sind, greifen 
kreuz und quer durcheinander. Die 7/andung des Vas deferens ist 
viel zarter als diejenige des Cirrus und lasst keine besondere 
IJuskulatur erkennen. 

Die Hden liegen als grosse, rundliche gebilde im 
Mitt elf eld des Gliedes zwisohen den Dotterstocken, den Keimstocke 
und dem vordern gliedrande. Genauere Angaben uber deren Zahl zu 
geben, ist wegen vorgeruckter Reife der Glieder unmaglich; dagegen 
sie ihres grossen Durchmessers (0,054 mm) v.egen kaun in grosserer 
Zhal auftreten. Bei Proteocephalus ocellatue , dessen Hodenblaschen 
beinahe gleichen Durchmesser (0,057 mm) besitzen, fand Kraemer (53) 
p -30 in jeder Prcglottids/^ 


It will "be noted that in practically every particular 
his description agrees with my figures 36 and with my description 
of the male organs of P. f alias, sp. which is from Coregonus f ero 
and is from Zschokke's collection. It is "beyond a doubt the same 
species that Zschokke and Kraemer described as Taen a ocellata 
and that Nufer in his table calls P. ocellatus . In the length of 
the cirrus-pouch in the shape and in the position of the con- 
striction, and in the bulbous inner end of the same, his description 
and my figure 35 agrpewell. The circular, swollen thickening 
which he notes in the posterior third of the cirrus can be best inte 
preted to be a coil in the ductus e jaculatorius. when such a coil 
is seen under certain conditions the appearance described by Hufer 
are simulated almost exactly Likewise in the position of the 
mass of coils of the vas deferens, in the number, size and arrange- 
ment of the testes nufer description and his tabulated data (P. 146) 
agree very well with my drawing and my description of the same 
organs in P. fallax n. sp. In not the slightes degree can his 
flescription of t e male organs of the species in question, he 
said to agree with my fig. 2, a delineation of the oro£lottid of 
P. ,:a crocephalus no with G. Schneider's or my own description of 
that species. 

Unfortunately I have not had the opportunity of examin- 
ing llufer mateaail of this species nor any other specimens of 
cestodes from the eels of Lake Lucerne. I cannot therefore make 
a positive 


determination of his form. The evidence at hand permits his 
cestode material ..rom Anguilla to become confused with P. falax 
from Coregonus fera(2) that the Anguilla of the Lake Lucerne harbor 
P. fallax, (3) that the ripe proglottids described by nufer had 
been taken into the intestine of the eel with its food, perhaps 
Coregonus fera , living or dead. 

The first suppostion is always a possibility when one is 
working with much material. The eecond supposition does not give 
complete satisfaction because Nufer was unable to find the fifth 
ucker which aught to be easily found in specimens of_P. fall ax 

The third supposition permits Nufer* s entire specimens to 
be considered as P. Macro cephalus . It also permits the view that 
the ripe proglottidsi upon which Nufer apparently based his 
description of the proglottids became confused with the specimens 
of P. macrocephalus by natural means. 

The embryo of Nufer' s form is about the size of the 
embroy of P. macrocephalus as determined by me. Nufer 's measurement 
of the embryo was 0.0184 mm. while theerabryo of my specimens of P - 
macrocephalus measured 0.017- .019 and rarely 0.021 mm. in an 
ovoidal embryo. The embryo of__P. fallax is much larger, according 
to my measurements -0.0312-0.0336 mm. 

Thus for attention I have not been called to Nu^fer's 

! description of the pernale generative organs. He claims that the 
histological structure of the vaginae of P. ma_cro cephalus and P. 
ocellatus differs in some respects. My own investigations of the 

two species especial y ..hen the investigations of Kraemer are taken 


into consideration lead me to believe that the vaginae of the two 
species are alike in histological detail. 

Nufer described the ovary as beeing single, not bilobed, 
In this he ahs been led astray by a condition sometimes found in 
ripe end-proglottids. In such cases the two lobes of the ovary may 
be pressed backward until the ends of thelobes taouch in the median 
line. In my own investigations I have thus far failed to note any 
fusion of the lobes. ITufer's drawing of this condition is hardly 

ITufer also olaims the walls of the uterine pouches 
degenerate evenually so that the eggs come to lie in the parenkpma 
I have very much doubt the accuracy of thisstatemont . I have ex- 
mined many secions thru large numbers of old ripe proglottids of a 
considerable number of species of this genus and have failed to 
find any such degeneration of the uterine walls. Hufer's material 
must have been extremely poor condition, or else he wofully mis- 
interpreted the appearance of ths sections. 

My conclusions regarding ITufers form are: a) That the 
complete specimens taken by Nufer from Anguilla vulgaris were 
probably P. ma.CFocepb.alus. 

(b) That a part of the description was based on the two 
loose end-prop-lot ids which realy belonged to P. fallax and that 
these were ingested with the food of the eel or that they were 
accidentally put with the cestodes of Anguilla during the manipula- 
tion of the specimens. 


o. That the eggs measured by Nufer probably came from 
P. macroceohalus. 

Luhe (1909) gave but a very short diagnosis of this 
form but included no figures of it. 

SOURCE OF MATERIAL.- Some specimens which I have identified 
as P.macrocephalus were collected by Prof. H. B. Ward in the source 
Of a biological investigation of Sebago Lake, Me. during the 
summer of 1j07. He examined eleven specimens of Anguill a G^ysypa 
/^#f» Of these number five specimens had no cestode infraction 
whatever ; two yielded each two Proteocephalids ; one yielded one 
?roteocephalid and two together yielded eighteen cestodes and some 
pieces. If these eighteen cestodes eight were ProteocephalidJ, 
Some of the pieces also belonged to this group. Altogether then 
there were thirteen Protecephalids plus some pieces but there was 
no complete strobile, among them. The specimens described are from 
bottles No. 47, 54, 56, 7 l/2 of the Sebago Lake collection. 

EXTERNAL CHARACTERS.- The worms are long and slender. 
Ho complete strobila was present but a fragment from the middle 
region of a worm measured 120 mm. The maxium breadth observed was 
1.6 mm. The youngest proglottids are much broader than long 
measuring 0.33 mm. broad by 0.022 mm. long. Mature proglottid^ are 
broader than long measureing 1.0-1.20 mm. broad x 0.4u-0.63 mm. 
long. The length of ripe proglottids may exceed the breadth. Ripe 
proglottids measure from 0.90-2.08 mm. long by 0.491-1.20 mm. broad. 


The tiad is very short. It has a slightly rounded apex 
which is not marked "by grooves or furrows. The head is somewhat 
flattened dorsoventrally, being 0.320 mm. broad at the base of the 
suckers and about 0.150-0.160 mm. thick. 

The four almost spherical suckers have a maximum diameter 
of about 0.095-0.100 mm. with a sucker opening of 0.050-0.060 mm. 
The sucker have only a moderately heavy musculature. The cavity is 
fairly deep. 

A fifth sucker is not present. It is replaced by an 
endorgan such as was described by me for P. filaroides. This 
endorgan has no*\ elongated cells and not trace of muscles. A vey 
few cells of irregular shape are surrounded by a membrance which is 
homologous with a basal membrane. 

The enctorgan is situated few micro-millimeters below 
the cuticula and has no connection with the exterior. The organ 
measures 0.0.J0-0.025 mm. The endorgan in this species does not 
resemble that of £ercae for I have shown (P1.3 f lg$$7rf*) that 
P. £ereae possesses a muscular organ that is doubtless a functional 

The neck is long and broad and thin, freguently it 
measure from 0.225-0.300 mm. or more in breadth while its length 
varies from 5.0-70 mm. 


The stobilation is quite evident. The notches between 
the segments are deep and the angles of the proprlottids are some- 
what rounded. Many shallow longitudinal folds give the surface 
of the worm a roughened appearance. 

The genital aperture is marginal situated ah out the 
middle of the proglottid. Its position in the strobila alternates 
irregularly from side to side. There is no genital papilla. 

Interjial anatomy.- The excretory and nervous systems 
have not been carefully investigated* The musculature of the 
strobila is much as Benedict found it for P._ ambloplities tho the 
muscles are not as heavily developed. 

Male pjr^ans.- The teste^fig.2. ) lie in a single layer 
in the dorsal region of the medually parenchyma between the 
vitel] aria and anterior to the ovary. They a e closely packed to- 
| gether until many of them are more or less polygonal in shape. 
They measure 0.063-0.074 mm. long by 0.042-0.063 mm. broad and they 
number from 100 to 120. 

The vas deferens forms a thick mass of coils extending 
from the middle fof the proglottid to the very short cirrus-pouch. 
Schneider very well says that the mass of coils of the vas deferens 
in excentively on the side in which the cirrus-pouch is situated. 

The cirrus-pouch(p((f ig. ^ ) is very short reaching just a 
little way thru the vittellaria. Its length is about 0.160 mm. 
The ratio of its length to the proglottid breadth is 1/6 or even 
less. The cirrus-pouch is bent up toward the inner dorsal wall 
of the deurao muscular sac. 


Within the cirrus -pouch the ductus e jaculatorius forms but 
one or two coils before passing over into the short, but thick-walled 
cirrus. The cirrus is straight. I have not seen it protruded. 

^Female organs.- The vagina lies anterior to the cirrus- 
pouch, but the vaginal opening into the genital atrium almost dorsal 
to the cirrus pouch^/(,f ig$.Jf; * ) Very near the opening of the vagina 
is a weak sphincter muscle. The lumen of vagina in its first region 
is quite large. This however soon constricts and the remainder of 
the vagina is of small diameter . Pigs. H show^how the 

vagina opening anterior and dorsal to circus-pouchy toward the ventral 
side and then back toward the ovaries. Several species of Proteo- 
cephalus, P. jaercae. P. pinguis, P. exiguus, show this twisting of 
the vagina part way ar^nd the cirrus-po/uch. This point has not 
been investigated in many of the species but it seems probable that 
it is quite a constanc't character in the genus. 

The broadened region of the vagina bears a ciliated 
ining on it inner surface. The presence of the cilia could not 
be demonstrated throughout the length of the vagina. A small 
receptaculem seminis was found a little way anterior to the mid- 
piece of the ovary. 

The ovarye" (PI. / figf./i- ) is a heavy ' ilobed structure 
of much the same shape. As in P. percae andp £^2J^^ 
but shorter. 

The vitellaria are long follicular masses in the lateral 
fields of the proglottids. The folicles are closely compacted. 
An oocapt with heavy musculature is present, 


also an ootype with its surrounding shellgland. The ducts of the 
interovsri^ space bear much the same relations as in other species 
of Proteocephalus in which they have been investigated. 

The uterus^in ripe proglottids is made up of a median 
tube with 7-11-12-14 lateral outpocketings on either side. The 
septa between the uterine pouches are sometimes very thin and after 
a sfftuous course either cases quite thick* The uterine passage is 
a small tube which after a sinuous course into the uterus about the 
middle of the proglottid. 

The uterine opening are 1-2 in number. Schneider in his 
speicmens found only a a single uterine pore. In reality this 
difference between Schneiders finding and my own is slight and to 
inadeguate to serve as a basis for a specific distinction. A 
second uterine pore may be very readily overlooked. Especially is 
this time in some species in which the second pore when present is 
smaller than the first and is situated some distance from it. 

Eggs.- These were taken from the uterus of alcoholic 
specimens. The thin outer hyaline shell characteristic of the egp;s 
of many species could not be demonstrated. The outer shell in this 
case corresponding to the eecond shell ox most Prpceophalus species 
was thick and granular. It measured 0.026-0.036 mm. in diameter- 
0.026-0.029 mm. being the more common dimensions. The erabryoifc 
self, varies from spherical to ovoidal. The diameter varies from 
0.017-0.019 mm. and rarely 0.021 in a long oval embryo. 


RELATIONSHIPS.- The indenity of this species with 
Lin-cow's P . df&tatus has already "been discovered whether it be the 
sane as P. he n i s pnaer.gcuS cannot now he determined. A discussion 
o±" its relationship to that form is given under the description of 
P. heraisphaericus Americany It show no mailed resemblance 

to any other oi' the species of/Proeocepnalus. In some ways it 

shows marked resemblances to P. c srnae H. Sp. from Aoerina cernae 

7. — a— 

It differs from that species in lacking a fifth sucker, in having 


many more test which are smaller , in having a much shorter cirrus- 
pouch aojStially and relatively. In have a much more voluminous mass 
pr coils of vas deferens, in having smaller embryo in the eggs. The 
proportions or the progiottids are also different. 

7/hile in some respects this species resembles P. percae 
its lack of a fifth sucker readily differentiates thea^e two species 
moreover its very short cirrus-poucn is in marked contrast to the 
very long cirrus-pouch in P. percae . 

It is greatly different from P. tor^losus in size in the 


si.:e of cirrus-pouch in the number of uterine pouches. 

The key to the species of the ganus and the tables 
will best show its relationshipes and its position in the genus. 


(PI. / figs. 9-17) 
1900: Proteocephalus ^ilicollis Benedict, 1900: 355-365. 

SPECIFIC DIAGNOSIS:- Characters of genus. Strobila short 
and slender. Length 9-15-25~o8 mm. Max. breadth 0.425-0.646-0.8 mm. 
Se -mentation not evident. Intersegmental furrows shallow. Neck 
2-4-7-10(?) mm. long "by 0.10-0.20 mm. "broad. First proglottids 
longer than "broad or nearly guadrate. Mature and ripe proglottids 
longer than "broad. Ripe proglottids considerably larger than 
mature ones, 0.680-1.190 mm. long x 0.460-0.595 mm. "broad. End 
proglottid elongated, pointed posteriorly, 0.714-1.50 mm. long "by 
0.40-0.646 mm. braod. 

Head somewhat globular, flattened dorsoventually , bearing 
four suckers on its broadest part. Breadth of head 0.120-0.160- 
0.170 mm. Suckers 0.058 mm. Broad by 0.069-0.035 mm. long. Opening 
of suckers about 0.040 mm. in diameter. Fifth sucker, muscular, 
functional, 0.037-0.048 mm. in diameter. 

GEIIITAL ORGANS .- Common renital sinus alternating 
irregularly, situated near middle of lateral margin of proglottid. 

MALE 0RGA1TS . - Tests 35-54 in number, in one layer, between 
vitellaria anterior to ovary. Diameter of tests 0.04-0.05 mm. 
Vas deferens forming mass of coils in midfield, Cirrus-pouch 0.209 

0.346 mm. long, reaching to middle of segment. Cirrus-ptraight , 0.10 
mm. long when protruded. 


FEMALE 0RGA1TS.- Vagina anterior to cirrus-pouch, cro sing 
cirrus^-pouch near middle. Sphincter raginae weak, 0.008 mm. thick. 
Lobes of ovary retort-shaped with smooth outlines. Vitelline 
follicles small, not compact. Uterus when fully developed with 
9-14 lateral pouches on either side. 

EGGS.- Embryos about 0,019-0.21 mm. in diameter, second 
egg membrane 0.036-0.046 mm. outer membrane 0.038-0.060 mm. 

HABITAT: In stomach (probably by post-mortem wandering 
and intestine. 


Coregonus -nigripirmis 

> jo 

ro gnathus 




Lake Michigan, : H. B. Ward, 
near Charlevoix 
" " " " : H. B. Wafd. 

" : H. B. Ward. 


La Rue. 
La Rue. 
La Rue. 

TYPE.- Alcoholics ITo. Ch 7 e, Ch 2 e, and an unnumbered 
bottle, in Dr. H. B. Ward's collection. Prom Goregonus Ijigripinnis , 
C. ^rognathus, and C. ArtedJ (type hosts) , Lake Michigan near 
Charlevoix ( t|5g/e locality)' 


HISTORICAL SUMMARY.- Benedict (1900: 555-365) described 
some cestoes which had "been collected by Dr. H. B. Ward while engaged 
in a "biological investigation of Lake Michigan Fish Commission during 
the summer of 1894. The hosts were Goregonus niRripinnis . C. 
progn«|thus , < q-tedi . Benedict who had for comparison some 

specimens of P_j_ ^ all ax (the Taenia oeellata of Zschokke) which 
had been nearly dried out in transit thought his material and P. 
/aliax (T. ocellata of Zschokke) identical. Accepting #raemer's 
statement that T. /ilioollis Rud. Aft/ T_j_ baellata Rud. were identical 
he used the name P. / ilicollis (Rud) to designate his form. Later 
he discovered that Taenia ooellata Rud. had page-precedence in 
Budolphi's original (1803) paper and in a foot-note he said that the 
name P^ o jellata (miss-spelling for P. ocellatus ) should be (s)g/ed 
to designate the form. 

DISCUSSION.- Careful examination of prepared specimens 
from came of the same lots used by Benedict and examination of one 
of his slides, and a comparison of these v/ith specimens which 
Zschokke had sent to Dr. H. B. Ward under the name Taenia acellata 
(y\ow No. 09.0 in Professor Ward's collection) has convinced me that 
Benedicts species is new. I propose for it the name Proteocephalus 
exiguus b y reason of the small size of the individuals ^Benedict 
worked out the morphology of the form quite correctly and probably 
would have given due weight to differences between his own and 
Zschokke' s specimens had the latter been in good condition. 


MATER I AI • - This species is "based upon the work of 
Benedict" ( 1900 : 355-365) and upon a further examination of, and 
collection of data from, certain lots of. alcoholic material in Dr. 
H. B. Ward's collection from will eh Benedict secured his specimens 

or study. These bottles bear the labels "Long-jaw, stomach. 
Charlevoix Jy. 16, '94 "ch. 7 c, Cestodes, Etc. Stoiaach Black-fin 
Charlevoix, Jy. 16, '94. ' 

"Ch. 3 c, cestodes. Coregonus Artadi Jy. IE, '94" 
Sli es have been prepared from the first two lots. 

EXTERNAL CHARACTERS.- The specimens which I examined care 
fully ranged in length from 9-10-15 mm. Benedict found the length to 
vary from 12-16-25 and one specimen only attained the length of 
38 mm. Th's one case seemed to have been due to an extreme afterfta- 
tion for the neck was 7 mm. long. This atterHation may have been 
caused by the method of killing. The maximum breadth Varied from 
0.425-0.595-0.646 mm. while Benedict records a maximum of 0.8 mm. 

The head (PI. / figS'/A/*"] according to Benedict 5 a& 0,120 
mm. broad. I found it to be 0.148-0.160-0.170 mm. broad. It is glob 
ular, The suckers which are placed just above the braodest zone are 
directed slightly anterial. They are 0.069-0.085 mm. long by 0.058ram 
broad. The sucker opening is e. 042-0. 048 mm. x 0.027-0.032 mm. 


Benedict states that the sucker is 0,040 mm. in diameter, but it seem 
that he must mean the diameter of the opening and not the diameter 
over the whole sucker* 

A fifth sucker (PI. I figfc/M-0 is present, 0.037-0.048 mm. 
in diameter. This fcom toto preparations ia apparently a true sucker 
Benedict says it has the "same structure (PI./ lig./O and 
musculature that the four large suckers possess." 

The neck measures 2-4 mm. in length by 0.110-0.120 mm. 
broad. Benedict fount the neck to be 5-10 mm. long by 0.10-0.20 mm. 
broad. The neck possess over into the first proglottids almost in 
perceptible. The first proglottids are longer tha broad or nearly 
quadrate. These very minature p -oglottids are few in number. The 
anlage of the genital organs appear very early and in a few proglotti 
further back mature sex organs appear. These mature porglottids are 
longer than broad. Thejr are few in number. Ripe proglottids are 
longer than broad. Three of these measued 0.935 x 0.595 mm. 0.680 x 
0.560 mm. 1.190 x 0.460 mm. Ripe proglottids are considerably larger 
than matmze . The increase in length, breadth and thickness is 
rapid. The longest proglottid in the strobila is usually the last 
one. Sometimes also this is the broadest segment. There terminal 
proglottids measured 1.190 x 0.646 mm. 0.714 x 0.510 mm. 1.500 x 0.4oT 
mm. the length being stated first. 

The end-progl ottids of variable form is functional. It is 
nearly alwasy the largest proglottid. 


It usually tapers to a blunt point at its posterior end where the 
excretory pore is situated. 

The segmentation (PI./ figs. ft/0) is not evident. The 
angles at the ends of the proglottids are scarcely marked. The 
inters egmental furrows are shallow. 

The total number of proglottids in a strobila is small 
10-12-14-15-20. Benedict found one specimen with 40 segments. 

IHTEEUAI AHAT01TY.- The histological structure of this 
cestode I have not worked out. Benedict's findings on these points 
are probably quite correct and need no checking over. 

GENITAL OR GAITS . - The common genital sinus is situated n 
near the middle of the margin of the proglottid, usually slightly 
anterior thereto. It alternates irregularly: 

Male Organs . The testes (PI./ fig.// ) are from 37-50- 
54 in number, 35-50 according to Benedict. They measure 0040-0.045 
0.050 mm. in diameter. The testes lie in a single layer between the 
vitellaria and anterior to the ovary. 

In my toZ^o mounts I could not make out with certainty 
the coils of the vas-de erens yet they seemed to lie in the middle 
of the proglottid near the dorsal wall. Benedict figures the vas 
deferens as formaing a coil extending beyond the middle of the 

The cirrus-pouch (PI. / fig. /y ) is elongated, slightly 
broador near its inner extremity and slightly constructed near the 
middle. It reaches the middle of the segment or even beyond that 


point in mature progiot tids. Its length micreases with the maturity 
of the segment. The following measurements of its length were taken 
from successive proglottids, 0.289, 0.296 x 0.513, 0.330, 0.341 mm. 
the last "being the maximum length found. 

The unpr otruded cirrus is straight. I have not seen it 
evaginatcd. According to Benedict the evaginated cirrus extends. 
0.1C mm. from the male opening . The ductus ejaculatorius is 

Fenalo Organs.- The vagina (PI. / fig$.yv ) opens into 
the common, genital sinus anterior. In a single proglottid the 
vagina was found posterior to the cirrus-pouch. This is probably to 
be considered an abnormablity or a very rate variation. A weak 
sphincter vaginae occurs a short distance from the vaginal opening. 
This sphincter is 0.008 mm. thick and is circular in cross section 
(Benedict) As Benedict has described it, the vaginae, extends 
straight in for a short distance then ir'bends backward and in- 
ward pssing under the middle of the cirrus soUf." 

The ovary very much resembles that of P. ?allax ( Kraemer ' s 
Taenia pcellata ) in shape. The lobes are thick and solid. Benedicts 

comparison of their shape to that of a retort is very good. The 
ovary is early decadent. 

The relations of the organs of the interovarial space are 
typical of the genus as Benedict* has described them. 


The vitellaria are lossely falliculai. The fallicles 
are not large nor are they closely porked. They too are decadent 

The uterus (PI. | f ig^A'T) in ripe proglottids consists 
of a median tube and 9-14 lateral pouches on either side. These 
;ouches occupy the ventral part of the segment while the tests 
are dorsal. 

EGGS.- "The eggs which are found in the uterus have a 
diameter of 0.02 mm. "Benedict my measurements garae a range of 
0.019-0.021 mm., the emhryo and 0.036-0.046 mm. for the second 
membrane. The puter membrane measures 0.038-0.060 mm. The second 
membrane is thick and granular. The outer shell is uterine eggs is 
thick and Ayaline. 

RELATIONSHIPS.- This species somewhat resembles P. />asil- 
lais word but is differentiated from that species b ; the smaller 
size of its head, by its very much smaller suckers and smaller sucker 
openings. This species has a much longer neck than has P. pisi lus 
The character of the segmentation of the two worms is utterly 
different. In P. fasillus it is very evident while the construction 
between proglottids being very deep in P. e.-cigims it is not 
evident. The number of proglottids in P. / t> isillus is greater than 
in P. gxiguus . The ceils of" the vas deferens i n P. £ Uxsillue is 
much smaller than in P. exiguus end it lies almost wholly anterior 
to the cirru.. -pouch while in P. g -^iguus it lies more nearly \at the 
end of the 


cirrus-pouch. She testa are more numerous in P. gaeillus 
*nd the cirrus-pouch is much ahcrtes than in P. Exigums. The ovaries 
j n P« ^ -is illus have a very peculiar droop e,e, the lobes are bent 
, -trongly posteriad while such a condition is never so marked. i n P. 
s dymus . The porportions of the proglottids are greatly different. 

A study of the tahles and of the key will "best show 
their points of difference. 

P. g .ciguus . closely resembles the P. ^allax n. sp. 
It differs from that species in having a smaller head, smaller, 
■ackers a smaller fifth sucker a shorter neck, fewer proglottids, 
smaller proglot.ids, more tests, a shorter cirrus, when evaginated 
a greater number of lateral uterine outpocketings and differnnt 
propotions of the proglottids. The embryos of P. exiguus are smaller 
than in P. ^ allax. The differences are of such a character and of 
such a conLJoney as to constitute, speci Vic differences. 

This species resembles P. 4-gonis Barbieri even more than 
P. gallox except that P. a . ;onis t has not well developed fifth 
E .cker in total length and in the size of the proglottids. P. 
g xiguus is the larger. P. g ^ig uus has" more uterine pouches, fewer 
and larger teste than has l J . a donis . Unfortunately Barbienis 
description fails in some particulars, nevertheless the sufficient 
data is given to establish the differences between the two species. 

P^e.ciguus resembles P. /.ibius gpe. but its head is 
smaller its sucker small ei , smaller proglot.ids, and a smaller 
cirrus-pouch. The e -tow species have about the same nurber of tests 
The embryo of P. 4 .bius is i"iuch larger than that of a?. e:iguus. 


P. exiffuuus differs radically from P. «.n'bijyiiu3 and P. 
gsoois in the possession of a fifth sucker. Other points of 
difference are also to "be found. 

P. e xifiuus has "been already compared with P. asillus 
word. It is smaller than other H. American species of Protecocephalus 


(PI. Pigs.) 

1887: Taenia ambl cpli tgs Leidy, 1887:22-23 

1887: Taenia micropterj^ Leidy, 1887:23 

1896: I cht h yo taenia ambloplitss Riggerbac$, 1896:267-268. 

1897: Taen ia ocellata Linton, 1897. 

1900: Proteoc ephalus ambl oplitas B enedict, 1900:339-355. 

SPECIFIC DIAGNOSIS:- Characters of the genus. Cestcdes 
large, 280-410 mm. long, 2.-2.5 mm. in max. breadth. Surface of 
body rough, with transverse and longitudinal furrows. Scolex 
prominent, 0.57-0.88 mm. broad, divided into quadrants by deep 
grooves. At tip of scolex is a minrte depression with a slight 
elevation in the center. Suckers large, round or cvo.l in outline, 
deep, directed outward and anteriad. Length of suckers 0.300-0.400 
mm. Fifth sucker ncne, replaced by end-organ in tissue. 

Prcglottids closel;, joined together, corners rounded. 
Young proglottids 12-15 times broader than long. Mature and ripe 
proglottids broader than long, about quadrate, or rarely longer 
than broad. 

Genital or gans :- Genital sinus irregularly alternating, 
marginal, situated about l/4 the length of the proglottid from the 
anterior end. Vagina opening anterior to cirrus-pouch. Vaginal 
sphincter long and very thick. Receptaculum seminis present. 
Uterus with 15-20 lateral outpccket ings on either side. 

Cirrus-pouch pyrifcrm, musclar, 0.400)0.500-0.550 mm. 
long, reaching 2/7-2/5 across the proglottid breadth. Cirrus, when 


protruded, long and slender. Ductus e jaculatorius much coiled. 
Vas deferens forming a greet mass of coils extending to middle of 
proglottid or past. Testes 70-1CC, lying between vitellaria. 

EGC-S:- Outer shell ellipsoidal, 0.036-0.043 mm. in 
length; middle shell 0.022-0.024 mm. in diameter; embryo 0.0168- 
0.018 mm. in diameter. 


HABITAT : - Intestine 





Ambloplites rupostris 

IJicropterus Salmoi los 

. jnblopl i tes rupestris 

Lake George, H. Y. 

Jos. Leidy 
Jos. Leidy 

Leidy 1887 
Leidy 1887 ' 

J. W. Milftfer Linton 189 7 l 

Micrcp torus dolomiej S 
Licrootorus salmoides 
Uicrop^orus salmoides 
Ami a c ftlva 
Amia calva 

Lake St Claire, Mich H. B. Ward 

Walnut Lake, Mich. 
Pelican Lake, Minn 
Lake Erie 

T.L. HankinsGn La Rue 

E. G. Davis 
H. B. Ward 

Lake StClairo, Mich- H. B. Ward 

Benedict 190C 

La Rue 
La Rue 
La Rue 

1. Recorded as Taenia micropteri . in "body-cavity. 

2. Recorded as Taenia ocellata, Rud. 




HISTORICAL SUMMARY.- This species was first described by Leidy 
(1887:22-23). His specimens were secured from the stomach of Amblo- 
pli t e's rupest ris. Lake George, Slew York. His diagnosis follows: 
: '' Taenia Ambloplitis .- Head quadrate, spheroidal, consisting almost 
entirely of the four large spherical bothria, with the summit 
slightly prominent and conical or depressed and unarmed; neck very 
short or none; body compressed cylindrical, gradually widening from 
the head tc near the posterior part, where it slightly narrows to 
the end;, segments linear, becoming gradually longer and wider, and 
then more quadrate, all deeply and pretty regularly wrinkled into 
two or iicrt • ' .'i.?. I "■• t f; ?-:12 de^pl y anci prett_ c 
two or three anriuli. Genital apertures obscure. Length 8 to 12 
inches; in alcohol contracted to 3 l/2 to 5 inches; greatest width 
2 mm. 

Head 0.5 to 0.625 mm. long and 0.75 to 0.875 broad. 
Bothria 0.575 mm. diameter. Commencement of body 0.625 wide. Ante- 
rior segments 0.125 long, 0.625 wide; subsequently 0.375 long and 
1.5 to 1.875 wide; posterior segments 0.75 long by 1 mm. wide. 
(Drawing. ) 

A number of specimens from "one stomach of the Rock Bass 
Ambloplit es rupest ris. Lake George, Hew York. 

Tb/s species resembles the Taenia ccell ata Rudolphi of 
the Eurppean Perch, Perca f l uviat ilis , and perhaps is the same. 
(Drawing . ) 

Immediately following his report on Taenia amblcplit j s 
Leidy (1387:23) reported Taenia micropteri from the intestine of 


Micropterus Salmoides nigrican s Lake George, H. Y. in the following 
word s: 

"Taenia "ic ropt er i . - Head large, compressed spheroidal, 
with four subterminal spherical "bothria and a papilliform unarmed 
summit; neck none; "bod:/ obscurely segmented, and with nc obvious 
internal organs, posteriorly variably narrowed and obtusely rounded 
at the end. Length from half an inch to an inch, and about 1 aim. 
wide. Apparently a larval form; found in the body cavity of the 
Black Bass, Micrcpterus nigricans . Six worms, soft, white, and 
active. The longer ones of an inch would elongate to double the 
length, becoming proportionately narrower. The head, about 1 mm. 
or more in diameter, varied in length and breadth, according to 
contraction, sometimes one and sometimes the other being the larger. 


Lake George, IT. Y. 

This form is very probably the larval form of P. ambloplit is. 
In dimensions and in general character the two forms are very much 
alike, even to the papillilform unarmed summit of the head. The 
obscure segmentation was probably due to wrinkling, not to a true 
segmentation. Since it has been shown (see report of hosts) that 
P. ambloplite s is thus far the only adult form reported from 
Llicropterus do lomieu and M. Salmoides , and since P.amb loplit is is 
known to occur in Lake George , the only locality from which Taenia 
Micropt er i has been reported I am caused tc conclude that Taenia 
micropt eri is identical with (Taenia ) Prot eocepbalus a mbloplit is 
(Leidv) . Therefore the name Taeni a mic ropteri should be stricken 
from the list cf valid species because of the priority of the name 
P.amb loplit is (Leidy) . 


Riggenbach (1896:267-268) put this species in his list 
of Ichthyotaenia yet he considered it a doubtful form. 

Under the name of Taenia o cellata R ud. Linton (1897) 
described this species from Am bio pi it es r upestris . His data agree 
very well with the data of Leidy, Benedict, and myself. A difference 
in the length of the neck caused Linton to doubt the identity of his 
form with Leidy T .a mblopli t i s. This difference is easily to be ex- ' 
plained in forms so contractile and so wrinkled as these. Linton 
himself says, "My specimens agree with Doctor Leidy 1 s pretty well 
except in the character of the neck. In T . ambloplit i s the neck is 
described as "shcrt or none" in my specimens the neck is long." 

Again in concluding Linton writes, "This reference of 
these Taenia of ambloplit e s of the species T . ocellata is provisional 
only. I think, however, that there ca^ be little doubt but that my 
specimens are identic&l with T . ambloplite s Leidy. The apparent 
absence of neck in Leidy' s species may be ascribed to the presence 
of strong transverse wrinkles, due probably to the action of the 
preserving fluid." 

Benedict (1900:339-355) working in Dr. H . B. Ward's 

laboratory redescribed this species using Leidy' s specimens for 


comparison with material which Professor Ward had collected from 
dclcmiep while engaged in a biological investigation of Lake St. 
Clair in August, 1893. Benedict usrV the section method frvnd 
that despite some discrepancies in size his specimens and Leidy' S 
agreed in anatomical and histological details. 


SOURCE OF MATERIAL . - The description which follows is 

based partly on Benedict's (1900) description of this species and 

largely upon my own work on material from bottles, #947 and #1056 

I collected by T, L. Hankinson from the intestine of Micropterus 

salmoides in the course of a biological survey of Walnut Lake, Mich. 

summer of 1906. Other specimens examined and identified by me as 

belonging to this species are as follows: In unnumbered bottle, col] 

LaRue, labelled, "Black Bass, Large mouth, Pyloric Region, Pelicdn 

Lake Minn. 1909." Collected by E. G. Davis; in bottle 10152, 

labelled, "Ex dogfish, Sandusky 0. Jy. 7, 1905."; In bottle #2d 

labelled, "7/10/09 P(ut) in B(ay), Host, Ami a Cal va-Int est ine . " 

-In bottle 3 85 ,T 7/l2/01 ?(ut) in B(ayJ Araia calyaT Tnte stine . 

In bottle #295, " 6/4/01. P(ut) in B(ay) . Ami a , Stomach." Bottles 

Nos. 10.152, 2d, d5, 295 are in Dr. H.B. Ward's collection. These 

together with Fos. 9.47 and 1036 coll. Hankinson which were sent in 

for identification were placed at my disposal thru the kindness of 

Professor Ward. 

EXTERNAL CHARACTERS . - This, one of the larger fish 

Proteccephalids, is the largest species of Pr ot eccephalus known at 

present from tne fishes of North America. It may attain a length 

of 410 mm. and a maximum breadth of 2.0-2.5 mm. The body is quite 

thick and heavily muscled. 

J Jn preserved specimens the outline of the worm is quite 

rough, due to the furrows on the margins of the strobila. The 

surface of the worn is also rough. Longitudinal and transverse 

furrows frequent. Expecislly noticeable is a deep median ventral 

furrow. On account of these furrows the st rabilat.ion is mere or 




less indistinct. Likewise the posterior limit of the neck is fre- 
quently obscured by numerous transverse folds which give an appear- 
ance of segmentation. 

The bead^is large and prominent. Its breadth is 0.75- 
0.875 ami. according to Leidy, 0.S2-0.88 according to Benedict. The 
beads examined by me measured 0.57-0.60 mm. Benedict describes the 
shape of the head thus, "Prom the anterior face it presents a nearly 
square outline, with -a deep notch in the middle of each side, di- 
viding the surface into quarters. Each quarter contains a large 
sucker which is directed outward and upward. The apex of the head 
is a smooth, rounded prominence with a small depression in the top. 
5To hooks are present. A fairly good idea of the shape of the scolex 
can be obtained by placing two truncated pyramids base to base. 
One of the smaller bases will represent the beginning of the neck, 
the other the prominence, while on the slope just beneath this 
would lie the suckers. The notches seen in the anterior view of 
the scolex form furrows down the four sides of the hea.d, which 
gradually decrease in depth and vanish on the first few proglottides. " 

Frequently the head is mere globose than Benedict des- 
cribes it. The small apical papilla may not always be seen, but 
the furrows have always been present in the heads examined. 

The suckers which are deep and well muscled measured O.oO 
mm. in length with a sucker opening cf 0.150 mm. in specimens ex- 
amined by me. Leidy 1 s measurement of the suckers was 0.375, long 
and Benedict's has 0.32mm. broad x 0.40 mm. long. In suckers of 
this size differences in contraction stages will explain the 
variation in size. 


There is no fifth sucker. This organ is replaced by a 
peculiar end organ lying deep in the tissues of the head. 

The first proglottios are much broader than long, measur- 
ing 0.344 mm. "broad by 0. 022-0, 0<j8 mm. long. Mature and ripe 
proglottics are broader than long as a rule, but at times may be 
quadrate, or even longer than broad. The last conditio! is rare. 
Th.e maximum breadth of mature and ripe proglottids i3 1.5-2.0-2.5 
mm. while the maximum length of the same is from 0.5-1.0-2.0 nun. 

The genital sinus is marginal, irregularly alternating 
and is situated about one-fcurth from the anterior end of the 
proglottid . 

I^TERITAL ANATOMY.- The cuticula, the musculature, and 
the nervous system were carefully investigated by Benedict who found 
nothing especially remarkable about these structures. 

The excretory system differs from that of P. fas s at us (Rigg; 
and Corg llo: pthrium lobe sum Rigg, in having no definitely placed 
Foramina secundaria. At irregular intervals from either dorsal or 
ventral excretory vessel arise numerous branches which take a 
fairly direct course to the exterior.. These branches are of various 
sizes and the lumen of each undergoes considerable variation in size 
in its course. The duct thru the cuticula is very small and not 
beset with small bristles as Kraemer (1892) described for P.f ilicolj is 
PTo special musculature can be found in the course of these ducts 
to the exterior. The main longitudinal excretory vessels are four 
in number. They are situated just within the derwo -muscular sac. 
The ventral vessel has the larger lumen. It is further differentiated 



from the dorsal vessel by its very thin membranous lining. The 
dorsal vessel has a lining apparently made up of columnar epithelial 
cells. Rarely however the dorsal vessel is dilated to a size 
equalling that of the ventral vessel and in these cases the struc- 
ture of the walls of the two vessels appears to be identical. 

No transverse vessel connecting the ventral or dorsal 
vessels in the posterior part of the proglottid has been found. 

In the scole* there is an intricate plexus of excretory 
vessels. Many ducts opening to the exterior by small pores place 
the coils of this plexus in communication with the exterior. 

GETTITAL ORGANS.- The common genital sinus opens about one 
fourth of the length of the proglottid from the anterior end. It 
is irregularly alternate in position. A genital papilla is not 
present. The vagina opens into the sinus anterior to the cirrus- 
pouch. i 

Mai e organs: - Tne testes|iare spheroidal bodies which are 
sometimes rendered polyhedral by pressure of adjacent testes. They 
measure 0.050-0.065 mm. in diameter. They number from 75-100 and 
are crowded into one or more irregular layers in the space between 
the vitellaria and anterior to the ovaries. 

The vas deferens forms a large mass of coils beginning in 
the midfield of the proglottid and reaching to the c irn; s-pcuch. 
the major part of this mass of vas deferens is posterior to the 

h^ i s a 1 c 

of whic>* is attached by heavy muscle strands to the dermo-muscular 

The c irrus- pouchy i s a large pyrifcrm sack the larger end 


sac. The pouch is 0.40-0.50-0.650 mm. long by 0.1b0-0.230 mm. in 
max breadth. The ratio of its length to the proglottid breadth is 
2:7-2:5. The cirrus-pouch is heavily musculed with both longi- 
tudinal and circular muscles. 

The cirrus when protruded is long and slender, a little 
larger at the base than at the tip. When within the cirrus-pcuch 
the basal part of the cirrus and has a broad lumen. 

The ductus ejaculatorius forms an intricate mass of coils 
within the cirrus-pcuch. The ductus is slender. 

F emal e organs .- The vagina^ always opens anterior tc the 
cirrus pouch. Very near its opening the lumen widens considerable 
and the vagina takes its course in a smooth arc anteriad and then 
posteriad. In this region near the opening the va.gina is surrounded 
by a large and trick sphincter vaginae. This sphincter is about 
0.030 mm. thick and about 0.300 long. This prominent sphincter 
surrounding the rather large lumen of the- vagina renders the vagina 
a very striding feature of the proglottid. 

The lumen of the vagina enlarges considerably beyond the 
sphincter. This region of the vagina is weakly ciliated on its 
inner surface. There may be a few coils of the vagina anterior to 
the ovary. A receptaculum seminis is here present which is differ- 
entiated from the subsequent part of the vagina by its histological 
structure. The vagina discharges into the oviduct after making a 
few coils in the interovarial space. 

The ovafr is bilobed in the posterior part of the proglo- 
ttid. Each, lobe is thick and broad and somewhat clubshaped. The 


lobes extend cut tc the vitellaria. 

From the midpiece of the ovary arises a muscular organ 
the occapt or gulping organ. From the occapt arises the oviduct 
which after making one or more ceils is jcinea by the vagina. From 
this point the oviduct extends to the ootype. Just as it enters 
the ofitype the oviduct receives the common vitelline duct which has 
been found by the union of the paired vitelline ducts arising in 
the posterior part of the vit ellaria. 

The ootype is surrounded by the so-called shell -glands , 
the individual cells of which are long and clubshaped. The ootype 
discharges into a somewhat muscular duct known as the uterine 
passage. The uterine passage extends anteri<fcd passing the ovary on 
the dorsal side. It discharges into the uterus from the dorsal side 
of the same about 0.Q80-0. 110 mm. anterior tc the ovary. Benedict 
(1900) stated that there was no uterine passage in this species. 
However, an examination of his slides and reconstruction drawings 
convinced me that a uterine passage wes present in his slides as it 
was in my own . 

The vitellaria are lateral and follicular. In this 
species trey do not extend posterior to the lobes of the ovary nor 
do they parallel the posterior margin of the proglottid. The 
uterus in young proglottids is a median ventral tube from which 
later 15-20 lateral outpocket ings develop on either side. There 
are 1-2-3 preformed uterine pores. 

EGGS.- The uterine eggs are covered with three shells, 
an cuter thin hyaline, a middle thick and granular shell and an 


inner thin membrane e closely investing the embryo. The cuter shell 
which is not spheroidal but ellipsoidal is about 0.036-0.043 mm. 
long. ' The middle shell is spherical 0.022-0.024 mm. in diameter. 
The embryo measures 0.01©-0.017 mm. in diameter. 

RELATIONSHIPS . - The American forms which most resemble 
this species are P. Pen lexus n> sp and P. nemat osom a (Leidy) . The 
latter spec ies may be the ssme-as P. ambloplites. P. p erplexu s t 
however, is distinguished from P. ambloplite s by its smaller head, 
smaller suckers, thinner and smaller strobila, by the smaller size 
ana the different location of its sphincter vaginae, by the posterior 
prologation of its vitellaria, by its much smaller cirrus-pouch, by 
its fewer coils of ductus e jaculatorius , by its larger number of 
testes, and by the smaller size of the eggs. 

P. ■tmbloplit i s greatly resembles P. nemat osoma . It dif- 
fers from that species chiefly in the somewhat larger size of the 
head and suckers. 

P. amblop lit i_s is larger than most of the European species 
of Prot eo c epha lus from fish and differs from e large number of these 
In not possessing a fifth sucker. It resembles P. tofuloeus in the 
lack of a fifth sucker and in size. It may be easily differentiated 
from P. t crulcsus by reason of its larger head and suckers, by its 
sphincter vaginae which for size and length is unique in the genus, 
by its more numerous uterine pouches, and by the different arrange- 
ment of testes. 



SPECIFIC DIAGNOSIS. - Characters of the genus. Strobila 
Ion? and slender. Observed lergth up tc 170mm, probably up tc 250 
mm in complete individuals. Max. breadth up to 0.90-1.0 mm. 

Head small , 0.250-0.30 mm. bread by 0.20-0.22 mm. long. 
Head bearing at broadest part four large suckers. Apical region 
of head frequently prolonged into a rost ellum-like organ. Organ 
unarmed. Deep grooves between suckers. Suckers cf variable shape, 
always with pointed apex, with shallow cavity and thin muscular 
wall. Length cf suckers 0.130-0.170 mm. breadth of same 0.170- 
0.190 mm. No fifth sucker, no vestige of same. 

Neck slender, 2.0-3.0 mm. long by 0.1-0.2 mm. broad. 
First proglottids broader than long, 0.255 mm. broad by 0.017-0.020 
mm. long. Mature proglottids up to 0.85 mm. broad by 0.34-0,37 mm. 
long. Ripe proglottids longer than broad or cjuadrat e, . 680 mm 
broad x 0.90-1.00 mm. lorg. Old spent proglottids up to 2.0 mm. 
long by 0.4 mm. Broad. 

Segmentation evident . Posterior angles of proglottids 
slightly projecting. 

GENITAL ORGANS.- No genital papilla. Genital pore 
marginal, irregularly alternating, situated at end of first l/4-l/2 
of proglottid. 

Male organ s. - Testes numerous, 75-85 or 90, in a single 
layer occupying entire field between vitellaria anterior to ovary. 


Breadth of testes 0.04-0.06 mm, length of same 0.07-0.10 mm. Vas 
deferens a large mass of coils in the midfield. Ductus ejaculatcr- 
ius with few or no ceils. Cirrus slender, straight, muscular. 
Cirrus-pouch slender, -nearly straight, muscular, 0.185-0.20-0.265 
mm. long. Length of cirrus-pcuch 2 l/2-3 times into proglottid 

Female organs .- Vagina always anterior to cirrus-pouch, 
never crossing the latter. Beginning region of vagina narrow. 
This region about 0.1 mm. long, terminated by weak sphincter 
vaginal. Adjacent to narrow region a dilated portion. Vagina not 
ciliated in any part of length. Vesicula seminalis present. 

Vitellaria with large follicles, not paralleling posterior 
margin of proglottid. Uterus when fully developed with 20-25 
lateral outpocket ings on either side. Uterine pores ventral, 2-3-4 
in number. 

EGOS.- With three shells. Embryo 0.014-0.0156-0.0168 mm, 
Second shell 0.026-0.031 mm, cuter shell 0.027-0.033 mm. 

EXCRETOP.Y SYSTEM . - Four main lateral trunks. Ventral 
vessels large, dorsal vessels much smaller. j& 11 main vessels 
sinuous or spiral. Many secondary openings to exterior. 

HABITAT.- In intestine of Lepisost eus ftlatystomus (type 
host), Illinois river, Havana, Illinois (type locality). 

TYPE.- Alcoholics Ha 12 d, e; Ha 29 f; Ha 30t; Ha 87 a; 
Ha 88 a; Ha 101k; Ha 110c. Slides from Ha 30ij Material in Dr. H. 
B. Ward's collection. 


MATERIAL.- In a collection of endoparasit es secured "by 
Dr. H. B. Ward from fish caught in the Illinois river at Havana, 
Illinois, were some specimens of Prot eocephalu s somewhat resembling 
the species which I have designated as P. perplexus . On account of 
the peculiar form of the head and suckers of this species I propose 
for it the name Prot eocephalus sir.gluari s n. sp. 

In all, eight gars, Lepisosteus pl atyQt omus Raf. were 
examined. Of this number one only was uninfected with this Parasite. 
This gar was infected with P. perp lexus. The numbers of P. singular! s 
present in each of the seven infected hosts were: 2, 7, 26, 2, 5, 
4, 8. Two gars had a mixed infection of P. singularis a .nd P. 
perplexus . 

Slides have been prepared from some of this material. 
EXTERNAL CHARACTERS. - The longest piece measured 170 mm. 
Complete strabilos perhaps measure up to 2.50 mm. or more. Its 

maximum breadth is about 0.90-1.00 mm. The strcbila is slightly 

the latter species 
smaller than that of P. yterpl exu s yet it greatly resembles the 

outward appearance. The more slender neck and the smaller head are 
its more readily noted points of difference. 

The head (Pl.,figs. A ) is small, measuring from 0.250 to 
nearly 0.300 mm. in breadth by 0.20-0.22 in length. At its broad- 
est part it bears four large suckers which when well expanded cover 
up nearly the whole surface of the head except the apical region. 
This region which is usually flattened, bears on its middle portion 
a slender protuberance which bears no hooks. This is not a 


rostellum. At times this protuberance is greatly contracted, be- 
tween- the suckers are deep grooves which in many cases cause the 
head to appear almost lobate. The suckers themselves are more or 
less variable in shape but they always show the pointed apex. The 
cavity is shallow, the muscular wall thin. The suckers measure 
0.130-0.170 mm. long by 0.170-0.190 mm. broad. 

A fifth sucker is not present. A careful search thru 
three series of sections has failed to reveal any trace of an end- 

The neck is long and slender, 2.0-3.0 mm. to the first 
traces of segmentation. The breadth is 0.1-0.2 mm. usually much 
narrower than the head. The neck grades over almost imperceptibly 
into the first proglottids which are very difficult to discern * 

The first proglottids are 12-15 times broader than long, 
being about 0.255 mm. broad by 0.017-0.020 mm. long. As the pro- 
glottids mature thfey increase greatly in size measuring up to 0.85 mm 
bi^oed by 0.34-0.37 mm. long. Proglottids in which the uterus 
contains some egrs measure 0.51 mm. long x 0.867 mm. broad or are 
even quadrate, each dimension being about 0.85 mm. The ripest 
proglottids become longer than broad, measuring 0.680 mm. broad by 
0.900-1.00 mm. long and in rare cases of old spent proglottids the 
length may be about 2.0 mm and the breadth about 0.4 mm. 

The segmentation is evident. Ho longitudinal furrows 
are present. Transverse furrows occur only at the division lines 
between the segments. The pcstericr angles of the proglottids 


project slightly beyond the anterior margin of the proglottid 

not "been thoroly investigated as to its histological structure. 
However, certain points have been worked out with some care. 

The cuticula, the subcuticula, the parenchyma, the 
muscles of the strobila and the nervous system were not seen to 
differ essentially from similar structures in P. ambloplitis. 

Musc I es of head . - The muscles of the head were worked 
out in series of transverse and frontal sections. The muscles of 
the apical protuberance of the head could not be worked cut with 
care but it cculd be seen to be well supplied with muscles expecialljf 
of the subcuticular layer. The peripheral parts of the apical region 
of the head contain many heavy muscles going up toward the tip. 

In the middle and lower region of the suckers 0.135 mm. 
from the tip of the head heavy muscles cross from one sucker wall 
to the wall of the sucker opposite forming a heavy muscle cross 
(P1.5 ( fig.9. Weaker musles cross the head dorsc-vent rally and 
laterally. These muscles extend to the subcuticular region and 
serve by their contraction to cause in part the deep grooves be- 
tween the suckers. Near the upper limits of the suckers at a depth 
of ebout 0.10 mm. from the tip of the head heavy muscles (Pl^fig^) 
connect the sucker wall with the lateral or dorsc-vent ral surfaces 
of the head. These probably serve a double function of causing by 
their contraction the deer groove between the suckers and of pull- 
ing the adjacent margins of the suckers together, thus broadly 


flaring the sucker. In this region the muscle-cress is weakly de- 

In the lower part of the head (Pl£figi$ just as the suckers 
disappear from transverse sections the fibers of the muscle-cross 
flare out very hroadly to secure an attachment on the sucker wall. 
In fact a great many fibers dip down below and attack themselves 
near the lower cuter margin of the sucker. At the sides of eac 1 
sucker large groups of very heavy muscles may be found the attach- 
ments and courses of which are best made cut in longitudinal sec- 
tions. Transverse fibers i.e. lateral and dorscvent ral , pass thru 
the muscle - cross and assist in the formation of a muscle-star 
(Plffig!*) Below this point the muscles rapidly assume the relations 
of the muscles in the strobila. 

The longitudinal muscles of the head are fairly well 
worked out. Certain groups which were of undert ermined character 
in transverse sections can be readily distinguished in frontal 
sections. The large groups of tangential! y cut ends ( Pl&f i g'JJ at 
either side of the sucker are in reality longitudinal muscles which 
come up from the neck region and find their attachment on either 
side of the sucker, right and left. Much weaker muscles bands 
pass up toward the tip of the scolex. In the apical region (Pl.^~ 
fig 3D many heavy muscles attached to the sucker wall pass out by 
diverging paths to the subcuticular area. These probably control 
not only a certain part of the movement of the suckers but also the 
form of the apex. 


The muscles of the head, both longitudinal and transverse, 
are much more strongly developed than in the head of P. f ilaroides, 
which I have worked out with great care. Their arrangement in the 
two forms is very similar. 

EXCRETORY SYSTEM.- In the head the excretory vessel may 
he followed as coiling and anastomosing trunks and vessels which 
are much less developed than in P. ambl oplitis or P. perplexu s. 
In the neck region the anastomoses of excretory system are much 
like those in the head. Here four main lateral vessels are to be 
found with branches end a very few openings to the exterior. 

A little further posteriad the excretory vessels in- 
crease in size. This is particularly true Of the ventral vessel. 
Both ventral and dorsal vessels have a very sinuous or spiral course 
even in the most elongated prcglcttids. In frontal sections (Pl.iT 
figtp the sinuous ventral vessels appear a.s series of oval or 
circular spaces in the parenchyma. In either frontal or transverse 
sections the ventral vessel is very prominent on account of its 
very large lumen. At frequent intervals branches may be seen to 
be given off from the ventral vessel and a lesser number from the 
dorsal. Many of the ventral vessel branches communicate to the 
exterior by mean? of small ducts thru the cuticula. These branches 
are more numerous and mere prominent than in any other Protec- 
cephalid examined by me. One branch opens quite regularly at the 
posterior angle of the proglottid. Other branches (by far the 
largest number) open on the ventral surface. A few branches of 


the ventral vessel open on the dorsal surface (P1.5~figIJ|)» 

No duct connecting the ventral excretory ducts in the 
posterior part of the proglottid has been observed. Nor have the 
relations of the excretory vessels been observed in the end- 
proglot t id . 

GENITAL ORGANS.- There is no genital papilla in which 
the genital pore is situated. The genital pore is marginal, ir- 
regularly alternating and situated at the end of the first third 
of the proglottid. Vagina and cirrus open into the common genital 
sinus, the vagina, always anterior to- the cirrus. In all the 
general features of the sexual organs these are typical of the 

Male organs . - The testes are oveidal or spheroidal in 
ihape, 0.04-0.06 mm. broad x 0.07-0.10 mm long. They are arranged 
(Pl.f,fig^ A ) in a single layer which fills the dorsal region between 
the vitellaria anterior to the ovaries. The region in which the 
coils of the vas deferens lies is free from them. They are 75 to 
850-90 in number. 

The vasa efferentia can be very readily made out just 
below the layer cf longitudinal muscles cf the dorsal side. These 
empty into the vas deferens which makes a large mass of coils in 
the middle of the proglottid and reaching over to the cirrus-pouch. 
This knot is relatively thick for it extends from dorsal to ventral 
wall of the demo-muscular sac. Entering the cirrus-pouch the vas 
differens or ductus ejaculations passes over almost immediately into 


the straight cirrus. The ductus ejaculations usually straight but 
sometimes it is thrown intc from one to three small coils. 

The cirrus itself has not been seen protruded. It is 
always slender, straight, and muscular. The cirrus-pouch (Pl^figtO 
is slender nearly straight, and quite muscular. It measures 0.185- 
0.200-0.265 mm. in length. Its length goes 2 1/2-3 times into the 
proglottid width. 

Female organs.- The vagina (P1.5"fig?tf in its first part 
has a narrow lumen. This region extends about 0.100-0.110 mm. 
from the vaginal opening. It is characterised by its rich covering 
of gland cell? and its weak longitudinal and circular muscles. The 
varinal sphincter which is poorly developed is situated just at the 
end of this first region. The sphincter is about 0.025 mm. long. 
Immediately following this region with the narrow lumen is a dilated 
portion which may extend clear to the vesicula seminal is. There is 
no ciliated region of the vagina. 

The vagina does not cross the cirrus-pouch but lies an- 
terior thereto until it comes to the knot of the vas deferens which 
it passes ventrally. It forms no coils anterior to the ovary but 
may be somewhat sinuous. 

The vitellaria are lateral. The fcllicles are large, and 
closely packed together. In the posterior part of the segment the 
vitellaria never extend posterior to theovary as in P. ^erplexus. 
The vitelline ducts are as in P. amblcplit is . 

Tbe lobes of the ovary are smaller than in P. frerplexus 


or P. amhloplit i s. They are more "branched than in either of those 
species "but this branched condition cannot be determined except in 

The relations of the oocapt , ootype, oviduct, lower 
vagina, vitelline ducts, and uterine passage are typical of the 
eenus and need no explanation here. 

The uterus^ is a median tube in mature proglottids. In 
ripe proglottids the median tuhe has 20-25 lateral out pocket ings ■ 
on either side which take up the entire ventral field of the seg- 
ment hounded hy the anterior and posterior margins and "by the 
ventral excretory ducts. 

Two , three or four ventral uterine pores have been ob- 
served, tho in most of the very eld proglottids the ventral body wall 
is split from end to end. m he foundation of the ventral unterine 
openings is as I have described it in P. fil oroides (La /fue 1910) 

EGGS.- The eggs very much resemble those of P.p erplexus 
in size hut differ in having a thinner and more hyaline middle shell 
which is never split into two layers. The embryo measures 0.014- 
0; 01 56-0. 01 68 mm. The second shell measures 0.026-0.031 mm. and 
the other hyaline shell 0.027-0.033 mm. 

RELATIONSHIPS.- This species in the relations of the 
head stands quite hy itself. Its proglottids somewhat resemble 
those of P.p erple x us bu t are smaller. In totib preparation the ventr- 
al excretory ducts of P. singular is "being much larger make a fairly 
easy means of separation. In the nurnher of uterine pouches the 


two species are much alike, but these pouches in P. perplexus ex- 
tend farther lateralad than in P. gingulari s. In the position of 
the voginal sphincter the two species are much alike hut the he- 
ginning region of one is dilated, in the other contracted. One 
has a ciliated vagina, the other not. The cirrus-pouch of P. 
singul ari s is more slender, the cirrus and ductus ejaculat orius are 
straighter anymore slender than in P. perplexus. 


(PI. 3 Figs. 3?-^ 
1886: Taenia /ilicolis Leidy i88b:62-3. 

SPECIFIC DIAGNOSIS.- The characters of the genus. 
Oberved length up "Go lb. 5 eras. Max breadth 1.7 ram. Strobilation 
evident, angles of proglottids sharp and distinct. First 
progiotuias rauen oroad-^r tnan iong. Mature proglottids broader 
than long 1.70 mm. broad x 0. 595 mm. long. P.ipe proglottids 
quadrate or longer than broad, l.u2 mm. broad x 1.10 mm. long to 
1.75 mm. long x 0.510 mm. broad. 

Head somewhat spheroidal, flattened dorsoventually , 
divided r/rco four quadrants by grooves extending nearly to the 
apex. At the apex frequently a small papilla situated in a 
depre.s ion. Head 0.663-0.714 mm. broad x 0.425-0.510 mm. long. 

Suckors four, 0.340-0.459 mm. long x 0.255-0.272 mm. broa 
Cavity of sicker deep. Ho fifth sucker, no nostellum, no trace of 
end organ. 

Heck broad no. T;nicK:, .v.bouu 0,5 mm. long. 
GEHITAI 0RGAI7S.- Genital pore marginal, situated at and 
of first of proglottid, irregularly alternating. Ho genital 


Male Organs.- Cirrus- pouch elongated oval in shape, 0.30- 
0.344 mm. long, extending 1/3-1/4-1/5 across the proglottid. Ductu 
ejr.c-.-latorius with 1-3 coils. Cirrus when protruded spindle shaped 
$3i±6ker at base, 0.60 mm. long. Testoc 135-155 in number. In one 


layer oceuping dorsal field between vit llaria anterior to ovary, 
testes 0. 069x0.037 mm. 

Female Organs.- Vagina anterior to cirrus-pouch., never 
crossing same, beginning region dilated. Spincter vaginae at end 
of dilated region. Inner surface of vagina heavily ciliated. 
Vitellaria vol min s, follicles of same large, compacted. Portion 
of vitelltria paralleling posterior margin of proglottid. Ovorian 
lobes thic , heavy irregular. Uterus v/hen fully developed with 
20-25 narrow lateral pouches. Uterine pores 2-4 in number. 

EGGS.- Uterine eggs with three shells, inner and outer shel.i s 
thin, inner thick, gran .ular somtimes partially or wholly split 
into two layers. Embryo 0.^13-0.014 mm. x 0.014-0.0156 mm. , second 
shell 0.019-C.030 mm, outer shell 0.024-0.036 mm. in diameter. 

EXCRETORY SYSTEM. - Two pains of main lateral canals with 
frequent anastomoses to exterior. Anastomoses in neck and head 


HABITAT:- In intestine. 

Host : 


Collector : 


Amla. calva L. : 

: Illinois river 
• Havana, 111. 

: H. B. Ward • 

La Rue 

jtoia caiva L. 

: Uorth Carolina 

: Jos. leidy 

:1a Rue 

iCepisosteus platystomus 

: Illinois river 

: H. B. Ward 

:La Rue 

TYPE.- Alcoholics Uos. (?) Ha 10 a; Ha 10 e; Ha 29 a; 
Ha 30 h; Ha 77 a, b,c ; Ha 89 a; Ha 94 e, h, and slides of 
Ha 89 a and Ha 94 h, in Dr. H. B. Ward's Collection. 

From Amia calva L. . (type host) Illinois river, Havana 
111., (type locality). 


HISTORICAL Leidy ( 1886 ; 62-3) found some specimens 
of a cost ode in Am la calva from North Carolina. These specimens, 
he thought might be Taenia f ilico l lis Rud. His specimens, 
judging from his data»probably belong to the species perplexus . 
Since Leidy published no drawings of his specimens no positive 
determination can be made. That part of his report containing his 
data is here quoted: "The worms accorded with the description of 
Taenia f ilicolis . e.t0. 

The worms accorded vrith the description of the Taenia 
/ilicolis, infesting Sticklebacks, Gastcrosteus . and is probably 
the same species. They range from 1 1/2 to 3 inches long, gradually 
widening from the delicate thread-like neck to the posterior rounded 
extremity, where they measure from 1 to 1.5 mm. wide. The head is 
spheroidal, variably broader or longer, and about e.625 mm., with 
the summit slightly prominent and unarmed and with four hemispher- 
ical, lateral bothria 0.25 mm. in diameter. Ueck variable, when 
extended long and narrow and usually about half the width of the 
head. Anterior segments, transversely linear, about an eighth 
the length of the breadth, gradually becoming inverted saucer- 
shaped or scutellate, and about one-fourth the length of the breath 
Posterior segments more quadrate, slightly widening behind, about 
0.75 mm. long and from 1 to 1.5 mm. broad; last segment longest and 
rounded. Genital apertures marginal. ^ 

SOURCE OF MATERIAL.- The specimens upon which this species 
is based were collected at Havana, 111., June and July 1910, by 
Dr. H. 3. y/ard. The hosts Amia calva and Lepisosteus platystomus 
were caught in the Illinois river at that place. Four Amia were 


examined, three were infected with 30,10 and 15-20 specimens of P. 
perplexu s respectively. One was uninfected. Eight gars were ex- 
amined; three were infected with 6, 3 and 5 specimens of P . perpl esux 
respectively. Five of the gars were not infected with this species. 
In all the Ami a and in one of the gars the P. perpl exu s was not 
accomparied by any other species of Prot eocephalus. In two of the 
gars P. singularis n. sp. occurred "beside P. Perpl exus . Amia calva 
is the type host. 

Nothing is known regarding the seasonal distribution of 
the parasite and almost nothing regarding its geographical dis- 
t ribut ion. 

EXTERNAL CHARACTERS . - Judging from external appearances 
alone these specimens were very much like specimens of P . amblopl it es . 
Even after staining and mounting some pieces in toto they appeared 
to belonp to that species tho certain points of difference could 
be indistinctly made out. Examination of sections revealed certain 
characters by which the two species covld be differentiated. I 
propose therefore to name this species Prot eocephalus perplexus 
by reason of the difficulties attending its differentiation from its 
nearest congener. 

In size P. perplex us is somewhat smaller than P. ambloplit es ( 
Its strobila is shorter narrower and thinner. Its head is a little 
smaller than that of P. amblopl it is but is very similar in shape 
and it has four grooves as dees that species. Sometimes the minute 
papilla at the apex of the head thought to be characteristic of 


P. ambloplitis may "be seen. The suckers are large and in a position 
similar to that which they occupy in P. ambl oplit is . 

The head (Pl.^figljJ) is somewhat spheroidal. It is 
flattened dorsovent rally and is evenly rounded anteriorly. Four 
grooves extend from the base of the head nearly to the apex where 
there is frequently a small papilla in a shallow depression. The 
head is not as truncate as Benedict (1900) described the head of 
P. ambl o pliti s . It resembles very closely heads which I have ex- 
amined and drawn (PI? fig v ) of the latter species. 

Four heads of_P_. p erpl e xus measured 0.663 mm, broad x 
0.510 mm. long, 0.697 x 0.425 mm, 0.697 x 0.510 mm, 0.714 x 0.459 mm. 
The suckers measured 0.340 x 0.255 mm - 0.459 x 0.272 mm. There 
is no fifth sucker and no rostellum. Sections thru the apex of the 
head failed to reveal even a vestige of a fifth sucker or of an end- 
organ. In P. ambloplit es the end-organ is large. 

The suckers are deep and well muscled usually longer than 
broad. The sucker opening is directed outward and slightly forward. 

Ho complete strobila was found yet most of the longer 
pieces contained ripe prortottids. Six pieces with heads measured 
65, 91, 106, 105, 125 and 155 mm. respectively. The maximum breadth 
observed was 1.7 mm. 

The worm shows a very evident strobilat ion, the angles of 
the proglottids are sharp and well marked. The posterior border of 
one proglottid overlaps the anterior end of the one following. 
Segmentation begins about 0.5-0.6 mm. from the head. Here the 
segments are much broader than long. Gradually the length increases 



in proportion to the breadth until in nearly mature proglottids 
the dimensions may he 1.70 mm broad by 0.595 mm. long. Further down 
the strobila nearly ripe proglottids may be about quadrate 1.02 mm. 
broad x 1.10 mm. long. Ripe proglottids are longer than broad, 
1.75 mm. long x 0.510 mm. Broad. 

In all but ripe proglottids the genital pore is not 
prominent but in these long proglottids the pore is situated on an 
eminence. Moreover in this part of the strobila the posterior re- 
gion of the segment is much broader than the anterior or middle 
re-ion. The neck is 0.5-0.6 mm. long. It is broad and thick, not 
as broad, however, as the head. 

GENITAL ORGANS.- The genital pore is situated on the 
lateral margin at the end of the first one fourth - one-half of the 
proglottid. The pores alternate irregularly. In mature and nearly 
ripe proglottids there is no genital papilla but in elongated ripe 
proglottids s -genital promirJe-ncle is noted. 

Into the common genital sinus open both vagina and 
cirrus, the vagina always anterior to the cirrus. 

Male org&ns .-The male organs (Pi;:ifig7p much resemble the 

male organs of P. amhloplit is b ut here again certain differences 

exist between the two species. The cirrus-pouch in P. perplexus is 

not as voluminous nor as long as in P. ambloplitis. The cirrus- 

. — 

pouch measures 0.300-0.344 mm. long. Its length goes into the 
proglottid breadth 3-4-5 times depending somewhat upon the state of 
contraction of the proglottid. 


Within the cirrus-pouch are from 1-3 coils of ductus 
ejoculat crius, a much email er bumber than in ambloplit es . The 
coils of the vas deferens outside the cirrus-pouch are very nem- 
erous. They form in mature proglottids a thick compact mass 
extending from the cirrus-pouch to the middle of the segment. 

The cirrus itself is well muscled. When protruded it 


is a long spindle shaped organ slightly thicker near the "base than 
elsewhere. It is nearly 0.60 mm. long. 

The testes are very numerous; 135-155 in number. They 
are arranged in one layer, occupying the entire dorsal field 
anterior to the ovaries and as far laterad as the vitellaria, with 
the exception of the small region taken up by a portion of the 
cirrus-pouch and the vas deferens. The testes measure up tc 
0.069 mm . long by 0.037 mm. broad, their short axis lying par- 
allel to the long axis of the worm. 

Female organs . - The vagina (PI .2 figs^' in its first part 
is considerably dilated. Thrbuout the full length of this dilated 
region there is a weak circular musculature. At the end of the 
dilatation is a. strong sphincter vaginae 0.053 mm. long and 0.015 
mm. thick. P^ ambloplit is has a long and extremely heavy sphin- 
cter vaginae ih ile in P. singularis n. sp. the sphincter vaginae 
resembles that of P. perple xus. Throuout its length as far as the 
receptaculum seminis the inner surface of the vagina is heavily 
ciliat ed* 

The vagina does not cross the cirrus-pcuch. After reach- 
ing the mid-field of the segment the vagina takes a sinuous course 
posteriad to the intercvarial space. A small receptaculum seminis 


lies just anterior or posterior to the midpiece of the ovary. The 
vagina forms no coils anterior to the ovary. 

The vitellaria (PUfigs^ are voluminous, the follicles 
are of good size and closely packed together. They extend not 
only to the extreme posterior margin of the proglottid hut are 
bent around and parallel the posterior margin nearly to the mid- 
field. This is particularly noticeable in ripe proglottids. The 
paired vitelline ducts pass throu this posterior contimation of 
the vitellaria. 

The ovary is bilcbed. The lobes are thick and heavy. 
The outline of each lobe is somewhat irregular due to proterber- 
ames. In sections it is noted that the ovary is not a solid organ 
but is made up of smaller parts more or less fused together. 

An oocapt, ootype, shell gland and other organs usual] y 
found in the int er-ovarial space in Prot eocepholids are found here 
and in the relations characteristic of the genus. 

The uterus, a median tube in mature proglottids, is made 
up in ripening proglottids of a median tube and 20-25 narrow 
laterial pouches. These pouches occupy the entire ventral field 
between the vitellaria and the anterior and posterior proglottid 
limits. There are 2-4 ventral uterine pores. 

EGGS,- The uterine eggs have three membranes, an inner 
closely investing the embryo, a middle membrane, quite thick and 
granular and frequently partially or completely split into two 
layers, and an outer membrane thin and hyaline. The embryo 
measures 0.013-0.014 mm. x 0.014-0.0156 mm. It is usually some- 
what elongated but is at times spheroidal. The second membrane 
measures 0.019K>.030 mm. and the outer one 0. 024-0. 036in diameter. 



EXCKETOP.Y SYST3M:- The execretcry system is made up of 
four main lateral canals which traverse the length of the strobila. 
The tv/o ventral vessels are larger than the two dorsal ones. At 
; intervals small branches arise from the main vessels and lead to 


the exterior. "So transverse excretory commissure was obsewed. 
In the head and neck the anastomoses of the excretory system are 
very complex. The main trunks are nearly straight. 

RELATIONSHIPS.* This species is closely allied to P. 
amblopl i ti 3 yet it differs from it in size, in the lack of an end- 
organ, in the size and location of the vaginal sphincter, in the 
posterior prolongation of the vitellaria, in the smaller cirrus- 
pouch, in the fewer coils of ductus e jaculatorius within the 
cirrus-pouch and in the greater number of testes and, in the size 
of the eggs. In size P. perplexus somewhat resembles P. s ing - 
ularis. It also resembles P. singu laris in the shape and propor- 
tions of its proglottids and in the character of the vagina and 
vaginal sphincter. These tv/o species differ in the size and 
shape of the head, suckers, and neck. Tho nearly of the same size 
the strobila of P. perplex us is the larger. P. perplexus has more 
numerous testes, a larger cirrus-pouch and more numerous ceils of 
ductus ejaculatorius. The main excretory vessels of P. perplexus 
are straight but they are very sinuous in P. singula ris. 

P. perplexus differs from all the European species in 
the position of the sphincter vaginae and in the extension of the 
vitellaria al ong the posterior border of the proglottid. 



1909: Ichthyotaenia agonis Barbieri, 1909. 


SPECIFIC DIAGNOSIS, - Characters cf the genus. Cestcdes 
of small size. Strobila 3-4 cms long, max., breadth cf same about 0.5 
mm. Scolex small, 0.168 mm. in diameter, unarmed. No fifth sucker. 
Suckers circular, symmetrically arranged on head, size? Neck quite 
long, up to 3 mm. ) "breadth 0.140 mm. Neck not well differentiated 
! from strobila. 

Prcglottids very variable in form and number at times up 
to 50-70. First prcglottids broader than long, mature longer than 
bread, 0.580-0.620 mm. long x 0.330-0.370 mm. broad. Pipe prc- 
glottids filled with eggs, nearly quadrate, length and breadth about 
0.5 mm. 

Sex p rga n s . - Genital aperture marginal, irregularly alter 
nating, situated near middle cf proglottid length. 

Fem ale organs.- As in genus. Vagina always anterior to 
cirrus-pouch. Uterus in ripe prcglottids made up of 4-6-8 lateral 

Male organs.- As in genus. Testes spherical, 0.031-0.038 
mm. in diameter, about 100 in number. Testes packed in area be- 
tween vitellaria. Coils of vas deferens in middle of proglottid. 
Ductus ejaculat oris* nearly straight in cirrus-pouch. Cirrus 
straight, with a thick wall. Cirrus-pouch cvcidal, broad at inner 
end, extending to middle of proglottid. 


Ex c r e t o r y s y s t e m . - Two lateral excretory ducts rising 
in head in each lateral field. Excretory ducts united in last 
proglo'ttid to form an excretory vesicle. No secondary excretory 
openings in head hut present in proglottids. 

EGGS.- Diameter 0.037-0.038 mm. Barbieri failed to 
state whether this was the diameter of the entire egg or of the 
embryo . 

HABITAT.- In intestine and pyloric c&cca of Alosa f inta 
vSr. lacustri s Ea. Lake Ccmo, Italy. 

The material was collected by Prof. Ciro Barbieri at 
Bellagio on Lake Como in Italy from the intestines and pyloric 
caeca of Alo sa f inta var. lacustris Fa. He found them here in great 
numbers, up to 1200-1400 in a single host. Since I have net seen 
any of these specimens the description is based on the paper of 
Barbieri (1909). 

EXTERNAL CHARACTERS.- The scolex is very small, with a 
diameter of about 0.16S mm. It is unarmed and has no fifth sucker. 
The four suckers are circular in outline and they are symmetrically 
arranged on the head. 

The neck is not well differentiated from the strobila. It 
is about 3 mm. long and about 0.140 mm. broad. 

The proglottids are very variable in number and form. In 
one well developed individual which had ripe eggs in the last 
proglottid there were 50-70. The first proglottids are broader than 
long. Mature proglottids are longer than broad, 0.580-0.620 mm. 
long x 0.330-0.370 mm. broad. Ripe proglottids filled with eggs are 


• 4 


nearly quadrate, length and breadth being about 0.5 mm. 

The length of the worm is variable. Well developed indi- 
viduals measure 30 Tim. long while a single individual measured 40 

SEX ORGANS.- The genital aperture is marginal, irregularly 
alternating from side to sice of the strobila and situated about 
the middle of the proglottid length. It is marked by a shallow 
depression. In it the vagina and cirrus open. 

Female organ s . - The female organs are arranged as in 
other Prot eocephalus species. ?he ovary is bilobed and posterior, 
the lobes being oval in shape with smooth outlines. The vitellaria 
are lateral longitudinal follicular glands which lie just inside 
of the lateral exeretory ducts. Gl/iduct , ootype and other organs of 
the int erovarial space are as in other species of the genus. 

The vagina always opens anterior to the cirrus-pcuch. It 
widens slightly and again narrows in its course toward the middle 
of the proglottid. Near the inner end of the cirrus-pouch it bends 
pcsteriaS. i nd thence its course is directed to the interovarial 
space where it forms several coils. 

T^e uterus containing eggs is made up of lateral pouches 
(4-6-8 in drawing) which in fully ripened segments fill up the space 
within the vitellaria. 

Male organs . - The male organs are comprised of testes, 
the coils of the vas deferens, the cirrus, and the cirrus-pouch. 
The testes fill the whole area between the vitellaria and the 
ovar;-. They are nearly spherical 0.031-0.038 mm. in dic.meter. 


Barbieri figures about 100 testes. 

The coils of the vas deferens form a thick mass which 
occupies the middle part of the proglottid. The lumen of the vas 
deferens is large, its wall thin. Within the cirrus sheathe the 
ductus ejaculatorius has a sinuous scurse in the inner two thirds 
of the pouchy "but apparently it fcrrns no closely twisted ceils and 
then it passes over into the straight cirrus. The cirrus has a 
heavy wall . 

The cirrus-pouch extends to the diddle of the proglottid. 
It is ovoidal with its broadest part at the inner end. The wall 
of the pouch is thin, but thicker at the proxinal than at the distal 
end . 

The Excretory system .* The excretory system much re- 
sembles that of the other species of the genus. There are two 
lateral excretory ducts which take their origin in the head region. 
They empty into a small bladder situated at the base of the last 
proglottid. No secondary excretory opening'were observed in the 
head but. they were found in the posterior region of each proglottid. 

EGGS. - Diameter 0.037-0.038 mm. Barbieri failed to 
state whether this measurement included only the embryo or the 

egg and its membranes. 


Larval stages were found in Bythrot replies and $eptodoro . 
RELATIONSHIPS.- This species as will be seen from its 
position in the key and in the tables at the end of this report is 
most closely related to P. esocis (Gui. Schneider). It is differ- 


entiated frcm that species by its different proportions, by its much 
smaller and more numerous testes, by the position of the coils of 
the vas deferens ; and. by the relation of the vagina to the cirrus- 

P. agonl s while resembling P. am biguus in size differs 
from that species in having a longer cirrus-pouch, in the excentric 
position, of the coils of vas deferens, in having smaller testes 
which lie in one layer, in lacking genital papillae which that 
species has. 

P. agcnis is readily distinguished, from P. f alh&x , P. 
dub in s , P. pusillu s and P. exiguus by its lack of a fifth sucker, 
by its more numerous and smaller testes, and by its less nu/.iercus 
uterine pouches. 


PROTEOCEPHALUS 3ALM0ITIS-UMBLAE ( Monti cell i ) sp . in^ 

1813: Taenia salmon is omul Pallas 1813:409. 

1884: Taenia salmonis umble Sschokke, 1884:18-19. 

1891: Tetracotylus salmonis -umblae Montioelli, 1891. 

1891. Tetracotylus salmon!. -omul Ivlontieelli 1891. 

1896: Ichthyotaenia s almonis umblae Riggenbach, 1896:267. 

1896: Ic; th; : otaenia salmonis umblae Zschokke, 1896:783. 

1909: Ichthyotaenia salmonis luhe 1909:33 
HJoTllRIOkl SUMMARY.- This species was first described 
and delineated b v Zschokke (1884) from SalmOHvuM&alre Geneva. 

His description and firawing are inadequate for a detcrmi- 
nation of its position. Montieelli (1891 J considers this a species 
of Tetracotylu s. He thinks that Taenia salmonis Omul _ Pallas 
(Montieelli writes this with a hyphen) is a species dubia and 
considers it to be probably identical with T^ salmonis-umblae . 

Kraemer (1G92) does not discuss the species. Riggenbach 
(1896) does not discuss it from the standpoint of structure. He 
considers it a species Ichthyotaenia 

Zschokke (1896:783) lists this species in his sumnany 
of parasites found in about 400 fish from lake Geneva. He did not 
find it in any Rhein fish of which he examined about 1200. He did 
not write the name with a hyphen ^i^/as^ Montieelli (1891) and 
Riggenbach (1896). llufer (1905) did not report this species from 
the fish of Lake lucerne, luhe (1909) gives this species as one of 
the Ichthyotaenia. His diagnosis being short and concise is 



quoted in l«ein oi' Zeeh Okie's description (Zschokke 1884:18-19} : 

Ichthyotaenia salmonis umblae Zschokke, "30-50 c, • long 
1-2 mm. breit. am seheitol eine leichte sougnapf ahniiche Vertiefun^. 
Proglottiden longer als "breit, ea. 100 bis 150 an Zhal, Hoden 
blaschen Zaj3hlreich, ein hoydenfreies mittelfeld sehe^int zu 
fehlen, Cirrus -bent el bis fast zur mittelinie reiehend. Genitaloffnunj; 

ungefahr aTj\ der grenze vm 2 und 3. Funftel der Proglottidenlange. 
Uterus mit zahlreichen "(ca. 6?J Blindsacken jederseits. Ealkkorper- 
chen sehr zahlreieh, besonders in sooles und hals. 

Im darn von Salmo s alve linus 1. ; bisher nur aus dem 
Genfer See be kannt". 

DISCUSSIOII.- This species is too little known to enable 
me to determine its position in the genus. On account oi* its larger 
size and more numerous testes I am sure that is is not the same as 
£• fallox or P. duMus. These characters which separate this 
species from the two just mentioned cause it to resemble the form 
from Trutta fario which I have called P. negle ctus and it may be 
the same. For. the present the form must be considered as an species 
mquirenda. However, the name as Zschokke soelled thekcause it* is 
a trinomial , can have no standing. Therefore the hyphenated "tt****^ 
Monticelli wrote if must be retained for this species thus 
groteoeephalys salmonis-nnblae ( Monticelli ) . 

..nether this species is the came as Tetr acoty lus salmonis 
onu__ (Monticelli ) cannot be determined for neither species is 
sufficiently known to Justify a comparison. 


Rudolphi (18iy:7cJ says Taenia salmonis omul is probably a species 
Of 3 othrioce^hal$i s. Montioelli (1891 J calls it a species dubia. 
I myself feel certain that the species should be so considered. 


(PI. 3 Pip-5.5^5) 




0. P. Muller, 

1780: 152-155-179 



cyst ica 

Pallas , 

178.1, I : |of. 




Bat sch, 

1786: 234. 




0. P. Muller, 

1788, IT: 5. 




Schrank , 

1788: 48 





1790:3079 So. 77. 

? 1790: 




1790: 3079 No. 79. 




Eudolphi , 

180: 112. 


Hal y sis 


Zeder , 

1803: 355. 





1803: 376. 




Rudolphi , 

1810; 108. 




Rud ol phi , 

1819: 149. 








f ilicollis 

Gui . Schneider 

1902: 2] 


I chthyo taenia 


Gui Schneider, 

1902: 23. 




Gui Schneider, 

1903: 13-22. 




Gui . Schneider, 





Gui. Schneider, 

1905: 15-17. 


SPECIFIC DIAGNOSIS.- Characters of the genus. , Worms 
of varying length 20-200 mm Max. "breadth 1.1-1.5-2.0 mm. Head 
short, broad, flattened dorsovent rally , apex slightly elevated or 
flattened. At summit a fifth sucker, muscular, 0.033-0.040-0.060 
mm. in diameter. Pour suckers, muscular, with deep cavities, 
0.085-0.100-0.137 mm. in diameter. Suckers at broadest zone of 
head or immediately anterior thereto. Head 0.192-0.357 mm. broad, 
usually about 0.300 mm. Thickness of head 0.170-0.238 mm. Surface 
of head without furrows. 

Neck 0.170-0.50 mm. broad by 3.0-10.0 mm. long. Trans- 
ition to first proglottids impersept ible . 

First proglottids usually broader than long, 0.255-0.34 mm 
broad x 0.085-0.102 mm. long, rarely longer than broad. Mature 
and ripe proglottids broader than long, Breadth of mature pro- 
glottids 0.935-1.19-1.30 mm, length of same 0.255-0.340 mm. Breadth 
of ripe proglottids 1.10-1.7 mm; length of same 0.42-0.85 mm. End 
proglottids about quadrate. 

Proglottids few - 150 or more closely attached. Segmen- 
tation fairly evident, angles of proglottid rounded. Surface of 
worm wrinkled and rough. 

GENITAL ORGANS.- As in genus. Genital aperture marginal, 
near middle of proglottid, irregularly alternating. 

Male organs . - Testes in single layer, irregularly 
arranged between vitellaria. Testes ovoidal, 0.05-0.095 mm. long 
x 0.05-0.07 mm. bread, 50-60 in number. Vas deferens a thick 


straight mass of coils reaching to middle of proglottid or 
beyond. Cirrus-pouch long, slender, sying at right angles to 
margin of proglottid -- Length of cirrus-pouch 0.34-0.47 mm. long 
Ratio of length of cirrus-pouch to proglottid "breadth 1/3-2/5. 
Cirrus when protrvded 0.1-0.2 mm. long. Cirrus when unprotTu- ed 
straight within cirrus-pouch. 

F emal e or gan s.- Vagina always anterior and dorsal to 
cirrus-pouch. . Vagina crossing cirrus-pouch near middle. Vaginal 
sphincter small. Lumen of vagina ciliated. 

Ovary bilobed, posterior. Lobes long and heavy. Vitell- 
aria lateral, fallicular, denser near ovary. 

Uterus with 4-5-9 lateral outpocket ings on either side. 
Uterine pore single, situated near middle of proglottid. 

1GGS/'- Embryo 0.019, second membrane 0.0264-0.029, outer 
membrane 0.031-0.037 mm. 


j^ -Acerina cernua /. 


1 '?Cottus cixadricornis 


HABITAT .- Intestine of Percidae and others^, 


Perca n orvegica 

Perca fluviatilis 

Perca fluviatilis 

Perca fluviatilis 


Perca fluviatilis 

G-asterost eus 

aculeatus %. 

Perca fluviatilis 


Greif swald 
F inl and 



Bos sit en E. 
Prv. 3 si a 

Rossi ton E. 













0. P. Muller (1788) 
( Pallas )Rudolphi 1810 
(Pallas )Eudolphi 1810 
Rudolphi (1819) 
G.Schneider (1903*05) 

La Rue 
La Rue 

Miihling (1898) 
Muhling (1898) 
Lonnberg (1889) 

l). In the course of this work specimens from Aceri^ia 
cernua have been determined to belong to a new species. Outwardly 
these specimens much resemble P. percae and there is therefore some 
doubt whether these specimens collected hy Pallas and the Taenia 
percae Muller were identical. Such a doubt is valid until modern 
investigations show that Acerina cernua. harbors P. percae (Muller). 


2. Reeds confirmation. 

HISTORICAL SUMMARY , - This species was first described by 
0. F, Muller (1730-152-155, 179). His diagnosis (1780:179) reads 
thus: "TAENIA PERCAB capite bulbosc, ocellis quatuor, csculis 
marginelibus, afrticulis quandrangulis . " 

On account of its indcessibility I quote a portion of his 
observations (1780:152-3) tho in a measure these observations are 
recapitulated in Mullers (1788) description which I quote later (p. ) 

"Der Kepf Oder das erste Glied des vordern und schmalen 
Endes i st vor gleicher Breite mit den nachsten Gelenken, vorne 
stumpf , und in der Mitte des Randes gleichsam eingebogen. Oben mit 
zwei durchsieht igen und uber einander ste^henden kugel runden Knoten 
besetzt . Wenn der Kppf fast ausget rochnet ist, zeiget sich die 
Spur von vier Zirkeln, zwei an der Stelle der Knoten mit einem ein 
gedruckten dunkeln Punkte in der Mitte; und Zwei gleichsam im 
Schatten aa der untern Lef^e naher am Rande. 

Die Gelenke sind platter, und Kommen einem gleich Seitigen 
Quadrat naher, als be^den andern Bandwurmern. Aut beiden Seiten 
gegen den Rand laufen zwei hell e weiss e Lini en den ganzenn warm 
hindurch; eigentlich werder^ie, wenn man genau zusiehet, von den 
Verbindungen der Gelenhe unter brochen, und machen in jedem Gelenke 
eine kleine bogenf ormige Linie; hie und da siehet man neben den 
Verbindungen der Gelenke in der Mitte den gev/6hnlichen punktf orange 
Eindruck, und auf der einen Flache, nicht auf der andern, einen 


we is sen Q.uerst rj c v , der sich Von der Mitte des einen Seitenrands 
"bis zu^ker Mitte des Gelenkes erstrecket. Kie Gelenke sinfit am Re.nde 
dicker, als £ewohnlich; viele haben an dem einen aussern Seiten- 
Tarde ein tie#es Loch, und es scheinet dass der Querstricb von dem 
hier durc"K°f all erTd en Lichte herruhret, weil er an dem undurch- 
locherten Gelenke, und an der Seite, wo kein Lech, ist, nicht zu 
bemerken war. Uiese Locher sind wahre Vert ief ungen, und immer in 
der Mitte des Sei tenrandes ^er GelehKe, doch nicht in alien, 
sondern chne Ordnunp, bs,ld in zi?ei, bald in mehreren auf einander 
folgenden. Bei den andern Bandwurmeni habejich dergleichen nicht 
bemerket . M 


Under the name Taenlt cystica Pallas (1781, 1; 101) des- 
cribed what was said tc "be this species as a larval form encysted 
in the livers of the perch and the pike. Batsch (1736: 234) in a 
compilatory work referred to this species as Taenia percae. 

0. E. Muller (1788, H.: 5) gave a diagnosis and description 

which were accompanied by drawings. His work is sufficiently 


thorough as to permit a comparative study witfh^ suspect ed to he the 
same as his. His figures (1-4) have been faithfully redrawn and 
are reproduced on PI. figs. Eecause of its inaccessibility 

his diagnosis (1788) is here quoted verbatim: 

"TAENIA PERCAE . - Taenia caprte Bulbcso, coellis quatucr, 
osculis aarginalibus , articulis quadrangulis. Nye danske Vidensk. 
Selsk. Skrift. I, p. 64, 65, 69. Naturf orscher , 14 St. S. 152, 153, 

Taenia alternatim transverse lineata, arficulis quad- 
rangularibus , capite quadriverrucosc . GOEZE EI! lT GEWEIDEW. p. 416, 
t. 33, f. 11-14. 

Caput , seu exfremifas antica etfenuior inssquentium 
articulorum latitudine, obtusum supra bulbis binis sphaericis, 
ecularibus, pellucent ibus , subtus binis similibus o'bsoletis infra 
margins instructum. Art iculi corporis crassiores et planiores 
quam in conge£eribus figuram quadrangulam imitantur. In ipso cor- 
pore linea longitudinalis Candida utrinque conepicitur; haec propius 
inspecta e lineolis articulorum arcuatis quavis int ersect ione 

interruptis componitu.r. In ipsis intersect ioni bus porus seu os- 

*Cited from Pudolphi (1810) and Braun (1894-19C0) in Bronns Kl . 
tk» Ordnungen . 


culum sclitum passim conspicitur, ac m&ginis medio in altera, paginae 
parte, non in epposita, linea transversa alba ad medium paginae 
pertingens pellucet. Haec foramini in ipsissima ora laterali 
quorundam art iculc rum. ccrspicuc ac ad medium usque pertuso deberi 
videtur. Foramen hoc conaliculatum oviductus est ac in paucis 
Taeniae speciebus observatur. 

In paene exsiccato capitula quetuor circuli bulborum 
vestigia apparuere; posteriores puncto opaco impressi erant. 

In intestinis Percae nar ina.e; ultra viginti enim 
4iversa detate examini subieci, ir. duabus tantum Me,rtic et April'., 
unicam nempe in altera, in altera tres reperi. 

Quas Claris. G0S2E in Silurc reperit, sola absentia 
lireolarum lateralium non in omnibus deque visibilium differre 

Tab. XLlV.Fig. L Taeniam Percae natural! , 
Pig. 2 et 3. caput diverse ritu, 
Fig. 4. frustum articulatum corporis "inter- 
medium aucta magnitudine exhibent." 
Zool. Danica sen Animalium Daniae et Noruegiae Eaviovum 
ac minus notorum Descript i ones et Historia. 2:5. 
Drawings et Plate 44 figs 1-4. 


Ifuller (178B) gave Taenia alt e m at i m transverse lln eat a 
Goeze from Silurus glanis L. as a synonym of Taenia percae . 
Rudolph i (1810-108) recognized that Taenia alt ernatium transverse 
1 in eat a ^ceae was identical with Taenia o sculata Goeze and included 
it in his list of synonyms of the latter species. 

Schrank (1786), Gmelin (1790), Rudolphi (1801), and Zeder 
(1803) are inaccessible to me and these citations have been cited 
from Rudolphi (1810) and Brown (1894-1900) 

Rudolphi (1810) gave a diagnosis of the cestode from the 
perch under the name, Taenia o eel lat a, a name which he had previous- 
ly used for this form, Rudolphi (1801). It is especially to he 
noted that;, his synonmy Rudolphi gave Taenia percae Mull er as a 
synonym of Taenia pc ell a t a Rud. However since Muller's name Taenia 
percae is the earlier name and since it was suf f iciently well 
described and figured to permit of its redetermination that name 
should Ft and and the name Taenia o eel lata is to be considered only 
as a synonym and should be stricken from the list of Proteocephalus 
species. Rudolphi 1 s (1810) diagnosis, synonmy, and description are 
here quoted verbatum: 

Taenia ocellata. R. 

Taenia: c apit e hemisphaerico , csculis p reofu ndis, callo 
longii-isculc rugoso , a rticulis s ubquadratis l ineolatis. 

Pallas F. Kord. Beytr. 1 l.p.101. Tab. 3. fig. 33. sub 
Taenia cystica. 

Muller Naturf . 14. p. 152-154 et 179. T. percae. 

Ejus Zool. Ban. Vol. 2. p. 5. Tab. 44. fig. 1 - 4. t. percae. 


Batsch Bandw. p. 234. n.7 T. percae. 

Grael . Syst . H. p. 3079. n.77. (exclusa var./S) T. percae. 
ib. n. 79. T. cernuae. 

Schrank Verzeichn. p. 48. n.1^6. T. percae. 
Rudolph i in Wied. Arch. 111. l.p.112. T. ocellata. 
Zeder Naturg. p. 355. n.42. Halysis percae. p. 376. Hal. 
cernuae . 


Hab. In intestinis Percae f luvi at ilis et cgtrnuae Pallas 
copiose, Percae marinae Muller rarius, repererunt . Ipse vario 
anni tempore in Perca fluv. copiosam offendi. 

Bescr. Vermes a me reperti duos ad quinque pollices longi, 
postice lineam lati; Mulleriara icon octopollicarem sistit. 

Caput exiguurn, h.emisphaericum sub motu polymorphum, mox 
contractum, mox valde inflatum; oscula circularia, immarginata, 
excavata, valdeque profunda, cum Distcmatum piscinorum poris 
comparand^, valde mobilia, ocellcrum instar agitata. Collum satis 
longum, transverse rugosum, uti Corpus depressum. Hoc antrorsum 
gracilescit, postice subaequale, articulis anticis brevioribus, 
reliquis subquadratis, margine postico parum incumbent e , lateral! 
uterinque subrotundo. In utroque articulorum majorum margine 
linea lateralis, marginem neque anticum neque posticum omnino at- 
tigens, a.liaque transversa, vel a modii articuli parte in margin em 
decurrens, vel utrique margini laterali fere contigua. In 
speciminibus minimis lineae istae pellucidae desunt, eaedemque 
generationis negotio inservire videntur, uti obs. insqu. probabit . 

Obs. 1. Mullerus in articulorum majorum, non tamen 


omnium, margine altero (eodem) laterali foramen conspexit , quo 
linea" transversa (oviductus itaque) t erminabatur . Neque Pallas, 
neque ipse in verme saepissime a me axaminato, foramen istud 
observevimus , sed Mulleri specimina ma j era ideoque ad^tiora. fuere, 
neque dubitandum est, quin Percae marinae at fluviatilis vermes 
iidem sint uti et Taeniae a el. Pallas in Percae cernuae ventriculo 
et intestino repertae cl. virc non diversae visae. 

Obs. 2. Pallas Taenias a se in Esocis Lucii et Percae 
fluviatilis hydatidibus hepaticis repertas cum int est inalibus 
conjungit, et ipse in Percae hepate veraiem hydatide inclusum reperi, 

quern nulla prorsus ncta a Taenia ocellata distinquere valec. Cf. 

■ I 

Cysticercum ligutatum n.12. 

Obs. 3. Taenias a se in Perca cernua inventas per vigin- 
ti horas in spirits vini vixisse. Pallas auctor est. 

Obs. 4. De varietate /3. a. Gmelino perperam constituta, 
conf . obs. 3. in Taeniam osculatam n. 25. 


Rudolphi (1819:149) quoted his earlier (Rud.1810) short 
diagnosis. He further stated that Taenia o cell at a occurred abundant 
ly in Perca f luviatilis at various times of the year at Greifswald, 
that Pallas had found this species abundantly in the intestine of 
Perca cernua ; and that Muller had found it rarely in Perca norvegica 
a fish which Muller called Perca marina. 

Perca cerrua is no. known as Acerina cernua . Norvegica 
is Sebast es marina according to Jordon and Evermann (1896). 
Rudolphi (1819) stated however that Perca marina did not occur in 

the Muller* s region and that Muller 1 s specimens came from Perca 

h a 

norveg ica. It is doubtful if the species desiganted by Rudolphi 
as Perca norvegica and by Muller as Perca Marin a is identical with 
Sebas t er marir a because the latter is a marine fish whereas T aenia 
percae occurs only in fresh water fish. 

Bujardin (1845:583) and Diesing (1850:513) were content to 
quote the diagnosis of this species and its habitat as given by the 
earlier investigators. Von Linstow (1878:208,209) did not increase 
the list of host species. 

Zschokke (1884:16-17) reported and described a cestode 
from the Perca fluviatilis of Lake Lucerne which he identified as 
Taenia f ilicolli s Rud. His species is probably identical with 
Prot eOjSephalu s dubiu s n. sp. 

Under the name of Taenia ocellat a Zscokke (1884:13,14) 
described a ^estode from several hosts including Coregonus f era, 
it is not at al 1 unlikely that this species is the same as the one 


from Cregonus f era described by Kraemer (1892) first under the name 


of Taenia filicallis and then under the name of Taenia ocellata. 


^raemer's species has "been shown elsewhere m this monograph to be 
very different from Taeni a percae and it is in fact a new species 
Prct ecceph alu s falLefe , n . s p . 

Lonnberg (1889: 14) reported Taenia ocellat a Rud. from 
Perca f luviatili s at Upsala. His diagnosis is short yet sufficient 
data are given to enable one to determine that this form is not 
the same as the one reported by Lonnberg (1889:15) from Gjfst er ost eff s 
pungitius and identified by him as Taenia f ilicollis Rud. 
Lonnberg 1 s Taeni a ocella a is nearly 3 mm. braad by 150 mm. long. 
It is probable that this form is identical with Taenia percae , Muller 

Muhling (1898:36) found Ichthyot a enia ocel lat a in Perca 
f luviatilis and Gasterostaus aculeatus at Rossiten, E. Prussia. 
It is probable that his specimens from Gasteras^u s were Prot eoce- 
phalus Gast erost e i not P. perca e , and that his specimens from Perca 
f luviatilis were Proteocephalus percae . 

Guido Schneider (1902:21-22) reported a species of cestode 

in Perca f luviatilis which he considered to be Ichthyotaenia fi\)fi- 

callis . He believed this to be of the same species as that found 

by Zschokke (1884) in the same host species. He noted the fact 


that Kraemer (1892) considering his Taenia f ilicjallis to be a 
younger stage of Taenia ocellata put his two forms together in the 
same species, Tocellata . To this last Schneider did not agree but 
held that the forms from. Perca fluviatilis and Coregonus lavaretus 


were distinct species. He noted on the one hand the near relation 


ships and the hreat similarities of the species of the Ichthyotae nia 

and on the other the extreme variability of the cestodes. He 
concluded that it was not therefore wholly unlikely, that the one 
form stood in relation to the other as a variety, which has arrisen 
from the changed environmental conditions of the new host. He 
further gave a short description of the worm limited almost exclus- 
ively to external characters. 

In the same article Schneider (1902:23) reported some 
specimens of Ichthyotae nia oc el lata , which he found in Coregonus 
lavarat ius and also in Cottus quadrico^is . 

Schneider (1903:13-23) decided to avoid entirely the 
questions of identity surrounding the names Taenia ocellata ^ and 
Taenia f ilicolli s Rud. He proposed to use the old name Taenia 
, percae 0. P. Muller to designate that form from Perca fluviat ilis 
which he (1902: 21-23) had previously considered to he Taenia 
f ilicollis Rud. His reasons for so doing are here quoted in full. 


As synonyms of Prot eocephalus ( Ichth . ) percae . (O.F. Muller) 
Schneider gave Taenia percae 0. F. Muller 1788: 5 pi. XLIV Taenia 
ocellata Rudclphi, 1810: 108 . 

Taenia filicallis , Zschokke (part) 1884:165 
Ichthyotaenia ocella ta Riggenjaack (part) 1896:268. 
Ichthyotaenia f ill icall is Gui, Schneider (part) 1902: N 2, 21. 


It is to be noted that Schneider failed to include in his 
synonmy any part of Kraemer's (1892) Taenia ocell atus. This is 

the more remarkable when it is remembered that both Kraemer and 


Rigerenbach were students under Zscbckke and it/ quite likely t&at 
both Krae'ner and Riggenbach accepted Zschokke 1 s determination of 
that species, yet parts of Biggenbaehr* s and also of Zschokke' s 
data on P. f ilicolli s are held to apply to P. percae . Nor did 
Schneider refer to Benedict (1900) in his synonmy. 

In his later description of the species Schneider (1905) 
gave nothing more on the synonomy of P. percae but he added some 
descriptive data to that given in his papers of 1902 and 1903. 

Lii^e (1909) considered Taenia ocellata (Rud.) a synonym 

for P. percae (0. P. Muller). In his diagnosis he followed 

Draemer (1892) and as an illustration of the proglottid of the 
species he apparently redrew Kraemer's fig. 18. Apparently also 
he either ignored Schneider's work entirely or else he considered 
Kraemer's P. ocellatus and Schneider's P. percae identical without 

havinr .made a comparison of material. I have shown elsewhere in 


this work that Kraemer's P. ocellatus or P. f ilicollis is not the 


same as Schneider's P. percae . 

It is very different from Schneider's P. percae as a com- 
parison of the figures illustrating the two species show (see 
PI. 2 figs. ^and PI. 3 figs. ^3'^^ ). 

DISCUSSION.- A comparison of Muller's (1788) diagnosis, 
description and figures with Rudolphis (1810) diagnosis 

and description compel, me to "believe that the two authors were 
refering to the same form. A comparison of their data and Muller's 
figures with Schneider's description and figures, and especially 
with his material, is convincing that Schneider has found the 
same form that Miiller and Rudolphi descrihed as Taenia percae and 
T. ocellata respectively. 

Muller's fig. 1 (See copy PI. 5 fig. ? 3 ) compares 
very favorably with Schneider's figures and with his material. 
Muller's fig. 4. (PI. 3 fig. ? £ shows the shape of the 

proglottids and location of the genital organs very well. The 
light area extending nearly to the middle of the segment is 
doubtless the cirrus-pouch which under certain circumstances could 
have been observed by Muller. The heads are not unlike the heads 
of Schneider's specimens, which I have had for examinat ion. Taken 
all in all the identification seems to be as complete as is possible 
to make without a study of Muller's and Rudolphis actual specimens. 

Since the time of Rudolphi the species name$ P. f ilicolli ; 
and P. o cell, at us .h ave appeared frequently in the literature of 
Helminthology. Por the most part the names are listed in reports 


of parasites found, without diagnosis and without description. 
Since the appearance of Kraemer's work (1902) P. f ilicolli s and P. 
ocellatu s have for the most part been considered to be identical. 
Great confusion has resulted making it impossible at present to 
separate from the others the references to those specific names 
which should be considered in the synonomy and history of P. percae , 
An actual study of amny of the specimens hearing these names must 
be made before an exact determination is possible. 

A comparative study of specimens of this genus in the 
helminthological collections in Eurpoe would doubtless yield many 
interesting results. In the preparation of this monograph five 
lots of material identified as P. ocellatus Rud. have been studied. 
Pour of these lots were received from Europe investigators, the 
fifth from an American. Out of these five lots four have proved to 
be new species while the fifth proved to be identical with P. 
percae . 

MATERIAL.- Professor Ward secured from Professor 
Levander, Eelsingfors, Finland, some specimens of Gui . Schneider's 
species^ P p ercae 0. F. Muller and P. ocellatu s Rudolphi. This 
material included both alcoholics and slides. 

One bottle now "Mo. 10.123 in Professor Ward's collection 
bears the label; 

" Ichthyotaenia percae 0. F. M. 
Det . G. Schneider . " 

A second bottle now No. 10.122 in Professor Ward's col- 
lection is labeled 


"Ichthyotaeni a ocellat a Rud. 
Coregonus lavaretus . 
14, V, 02" 

The two slides were labelled " Ichthyotaenia ocellata , 
Coregonus, 24, VIII, 01." These must "be from the same lot which 
Schneider mentioned in his report (1902:23 and 53.) 

Sections and toto preparations were made from the No. 
10.123, P. percae, while sections only were prepared from the 
#10.122, Schneiders P. ocellatu s. Some of the heads from each 
lot were also studied in Glycerine. 

(See table ) of Schneider's descriptions of his P. 

ocellatus and his P. percae show hut two significant differences 
between the species. According to him the head of P. percae is 
much the larger. The main difference between them however is that 
the head of P. Ocellatus has "einen funften flachen Sftngnap^ von 
40yu im Durchwiesser an der Spitze, der offenbar auc^ noch funk- 
tioniert, da er aus einer grossen Anzahl von Radialmuskelf asern 
bestcht." While of P. percae he writes "An der cptize des Scolex 
findet sich das detttliche Scheit al-organ, welches bei konse^iert en 
Exemplaren eine kugelformige Anhaufung in der Lan^s^aXe des T/eres 
gestreckter Zellen darstallt, die ihren Charakter. al£ muskel-zelleU 
o^enbar verloren haben. Weir sehen hier ein rudimentares Organ, 
das, wie es scheint , jede Funktion eingebusst hat." 

All other differences besides this one can be explained as 
dur to contraction states. The breadth of the head when but a very 


few specimens can "be measured is a valuable tho not an absolutely 
reliable character in a form which is so variable and so contractu 
In measuring 1 Schneider's own specimens I found the heads of the two 
forms very similar. One head of P. percae was even smaller than 
the dimension recorded by him of P. ocellatus . while in every case 
observed by me the actual breadth of the head of P. ocellatus w as 
greater than that recorded by Schneider. The thickness of these 

heads measured by me was about equal to the hreadth that he re- 


corded. So this difference may be ruled out. 

for the fifth sucker, my examination of frontal 
section? of heads of P.. percae with the oil immersion lens has 

revealed the true muscular structure of a sucker. My drawing 

is ff/ 

shows the nucler, the radial muscles, the basement membrance , the 
out ends of the circular muscle fibers, and the cuticula continuous 
across the sucker. ITo cavity was noted in the fifth sucker of P. 
percae nor in Schneider's P. ocella tus. This last difference then 
between Schneider's P. ocellatus and his P. percae vanishes. 

An examination of the two lots of Schneider's specimens 
revealed a marked agreement between them in every diagnostic 
feature except in the proportions of the proglottids and this 
slight difference was undoubtedly due to different states of con- 
traction. LTost remarkable similarity was found in the relations 
of cirrus, cirrus-pouch, vas deferens, t est es, vaginia , vaginal 
sphincter, uterus, vitellaria, eggs and position of genital pore. 
An examination of the two comparative tables will show at once the 


similarities. The differences which Schneider noted in the sape 
of the ovary are readily explained as due to the contraction state 
of the proglottid. A comparison of drawings (PI. figs. and 

PI. figs. ) show the marked agreement between these two 

lots of material. 

Schneider's P. ocellatu s is clearly identical with his P. 
percae , and on account of the priority of the name P. percae (0.?. 
Muller) that name should be retained. 

EXTERNAL CHARACTERS.- These cestodes are of varying 
length from 20-200 mm. and the nax. breadth in the different spe- 
cimens varies from 1.1-1.5-2.0 mm. Differences in the stage of 
development . and also in the contraction states account for the 
greater part of this variation. , 

The short "broad head^is flattened dorsovent rally. The 
apex of the head may "be slightly elevated or it may he somewhat 
flatted. At the summit is a small but muscular fifth sucker. At 
the broadest zone of the head or just anterior thereto are situated 
four suckers which are directed forward and outward. The scolex 
varies in breadth from 0.192-0.357 mm. tho by f*r the greater 
number of heads measured were from 0.30-0.54 mm. broad. The thick- 
ness of the head is about 0.170-0.238 mm. such variations in 
breadth are due to the states of contraction of the muscles of the 
neck and head. Strong contractions of the longitudinal muscles of 
the neck and head cause the neck to dilate markedly. This con- 
dition also caused the posterior parts of the head to dilate. In 
such a state the suckers are directed more nearly forward. The 


converse of these statements also holds. With the relaxed 
longitudinal neck muscles and consequently the elongated neck is 
usually to he found the narrow head. 

The surface of the head is smooth without wrinkles or 
furrows. So also the surface of the neck is smooth. Sometimes 
transverse wrinkles on the neck simulate proglottids. 

The neck var/ies considerably in length and. breadth. The 
breadth varies from 0.170-0.50 mm. while the length varies from 
3.0-10.0 mm. The neck is thin. It passes over almost impercep- 
tibly into the first proglottids. 

The first proglottids are extremely variable in shape. 
In a strongly contracted worm they are much broader than long, 
0.34 mm. broad by 0.085 mm. long. In less contracted individuals 
these first proglottids may measure 0.255 mm. broad by 0.102 
mm. long. Schneider (1903) states that they are broader than long 
or, nearly quadrate or longer than broad. 

Mature and ripe proglottids show somewhat similar variation 
tho in less degree. Mature proglottids in all specimens examined 
by me were broader than long. The breadth var/ied from 0.935- 
1.19-1.30 mm. and the length from 0.255-0.340 mm. Fully ripe 
proglottids are slightly longer in proportion to breadth. They 
measure 1.7 mm. broad x 0.85 mm. long to 1.10 mm. broad x 0.42 mm. 
lonfr. Ho end proglottids were observed by me. Schneider describes 
them as about quadrate. 

The number of segments varies from the few in the very 


short worms to 150 or more in the longer specimens (Schneider). 
The segmentation is fairly evident especially when the worm is 
viewed with a 14ns of low magnification. The angles of the pro- 
glottids are rounded. The surface of the worm is more or less 

Y)€ O 

roughed oy shallow gro A ves and depressions. The prcglottids are 
closely attached to each other. 

The common genital aperture is situated very near or 
slightly anterior, to the middle of the margin of the proglottid. 
Its position alternates irregularly from side to side in the 
strobile . 


Mai e organ s.- The testes lie in a single layer between 
the vitella.ria and anterior to the ovary. They are of ovoidal or 
spheroidal shape. They measure 0.053-0.086-0.095 mm. long by 
0.05-0,07 mm. broad. Schneider's drawing of his P. ocellatus 
shows 50-6 testes the number which may be counted in preparations 
from either of Schneider's lots of material .were 0.10x0.06 mm. 


The vas deferens forms a thick straight mass of coils ex- 
tending from the end of the cirrus-pouch tc the middle of the 
proglottid or even a little beyond. In mature and ripe proglottids 
the vas deferens is always well aestended with spermatozoa. 

The cirrus-pouch is long and slender extending into the 
segment at right angles to the margin of proglottid. In a recon- 
struction (PI. 3 fig. (q 5 ) from trans-sections it is seen to 
curve upward, to the dorsal wall of the dermo-muscular sac where it 
is attached at its inner end by strong muscle fibers. Its length 


var^ies scmewhat in proglottids of different stages cf development 
and in different stages of contraction. It measures 0.34-0.37- 
0.425-0.47 mm. long, the greater' lengths occuring in greatly con- 
tracted, and hence very wide, proglottids. Schneider reported 
the ratio of its length to the proglottid hreadth as l/3 in his 
P. ocellatu s and l/3-l/2 in his P. perc ae . I find the ratio in 
each of his forms to he from 1/3-2/5. In no case did the cirrus- 
pouch reach to the middle of the proglottid tho at first sight the 
coils of the vas deferens frequently gave it that appearance. 

Schneider saw the protuded cjrrus in P. perc ae , where it 
measured 0.1-0.2 mm. In "both of his forms he found the cirrus 
straight in the sirrus pouch. The same condition were observed 
hy me 

"Fema le organs .- The vagina always opens dorsal to the 
cirrus-pcuch. This is shown hy a reconstructed trans-section 
(PI. } Pig. £tr ) thru this region. Then passing inward and 
bending slightly anteriad it crosses the cirrus-pcuch obliquely 
near the middle and passes toward the ventral wall of the demo- 
muscular sac. Prom the point of crossing the cirruspouch (PI. 3 
fig. ) it passes posteriad in a long curve to the interovarial 

space. The vagina lies just dorsal to the uterus. 

Very near the opening of the vagina a short sphincter 
vaginae 0.020-0.026 mm. thick may be seen. The inner surface of 
the vagina is apparently ciliated. A receptaculum seminis has not 
he en found. 


The ovary (PI. 3 figs.^* M * ,w ) is long and heavy. It is 
somewhat arched in the }ncre elongated proglottids and more elon- 
gated and slender in the longitudinally contracted proglottids. 
Frontal sections show the ovary to he made up of closely connected, 
tubes or branches. 

The vitellaria are long lateral follicular glands which 
extend from the anterior end of the segment to the ovary hut not 
beyond. The vitellaria are more dense near the ovary. 

The coils of the vagina, uterine passage, oviduct, and 
the common vitellaira duct nearly fill the int ercvaria.1 space. 
Here also are the occapt and obtype. The relations and connect- 
ions of these various passages are similar to those described by 


Benedict (1900) for this group. The uterine passage discharges 
into the uterus near the niddle of the segment. 

The ut erus^fig. L ^ ) in ripe proglottids has from 4 or 
5 to 7, 8 or even 9 lateral cutpocket ings on either side. These 
come to fill up nearly the whole ventral side of the segment. 
Schneider found but a single uterine pore near the middle of the 
proglottid. I have found this number to be correct. 

EGOS.- Schneider's measurements are as follows:- embryo 
0.025 mm. second shell 0.045-0.050 mm; outer shell 0.090-0.125 mm. 
My measurements are made from uterine eggs of alcoholic materials. 
I have already shown (La Eue 1910) that the outer shell swells 
up v/hen it comes in contact with the water. I am not sure that 
the inner one does this. Iffy measurements of his material are as 
follows:- Embryo only 0.019; second shell, 0.0264-0.029; outer 


shell, 0.031-C.037. Since I have had Schneider's own specimens 
for examination I am inclined to "believe that he measured the 
embryo plus its investing membrane, mhe eggs which I examined 
when so measured gave almost his figures for the diameter of the 
embryo . 

HOSTS.- In Perca f lu viat ilis, Coregonus la.varetus , 
possihly also in Acerina cernua. 

Locality.- Finland, Probably also in regions in the 
Baltic drainage basin, Sweden and Germany. 

RELATIONSHIPS . - Tho in some respects P. perca,e resembles 
P. agoni s , Barbieri P. exiguus n. sp. and P. pusillus , Ward, it is 
' much larger tv>an those species that the possibility of an identity 
is entirely prece4-e4ed. In many ways it resembles P. fall ax n. sp. 
but it is larger than that species. The character of the Proglottid 
is vastly different. P. percae has many more and larger testes, 
a logger ci rrus-pcuch , a much larger ovary, more voluminous 
vitellaria, a larger head, larger suckers, larger sucker openings. 
P. fallax has a larger number of preformed unterine pores. It 
also has much larger embryos. The shape of the region occupied 
by the ovaries in ripe proglottids is triangular in P. f allqx but 
elongated in P. percae . P. per cae resembles P. cernua e in the size 
of the head, in the size and character of the fifth sucker, in the 
shape of the ovary, and somewhat in the character of the proglottids 
P. percae however has in general larger suckers with larger sucker 
openings, a longer and more slender neck, a much longer cirrus- 


pouch, and fewer and smaller testes. The relationships of cirrus- 
pouch and vagina fire rreatly different in the two species. In P. 
percae the vagina crosses the cirrus-pcuch near the middle while 
in P_. cernuae the vagina does not cross the cirrus-pcuch. P. 
cernua e may have mere numerous uterine pouches. The embryos in P. 
ce rnua e are larger than in P. percae . 

P. percae resembles in some respects P. lcngicallis as 
described by von Linstow but it differs greatly in the number of 
uterine pouches, in the number of testes, the length of the cirrus- 
pouch and in the size of the eggs and size of the suckers. 

P. percae most nearly resembles P. pingufs n. sp. The 
heads of P. pingut s and P. percae are of about equal size. The 
suckers are much alike. The fifth sucker in P. pingues is much 
better developed than in P. percae . P p ingues is narrower and. 
regularly shorter. Many more of its proglottids are not as 
flattened as in P. percae . P pingues has many more uterine out- 
pecketings and uterine pores. Its cirrus-pouch is less than half 
the length of that of P. percae . 

P. percae readily differentiates itself from P. tomlosu s 
by reason of the total lack of a fifth sucker in the latter. In 
the number of uterine pcuches, in the length of cirrus-pouch and 
in many ether ways these species are very dissimilar. 



(pi. V figs.?*-?/) 

Taenia cyprini idi 

Vibor°: , 



Taenia articulis rotundis 




Taenia torulosa 




Taenia orbicularis 

Schrank , 



Taenia simplex 




Taenia torulosa 


17 90:3081 


Rhytelmint 1ms cyprini 

Zeder , 



Taenia torulosa 

Rudolphi , 

-± 7 


1803 : 

Halysis torulosa 




Taenia torulosa 

Rudolph! , 



Taenia Torulosa 

Rudolphi , 


1845 : 

Taenia torulosa 




Taenia torulosa 



1884 : 

Taenia torulosa 

Zschokke , 

1884:20, in part. 


Taenia torulosa 




Taenia torulosa 

v. uinstow 

, 1891:565. 

1892 : 

Taenia torulosa 

Xraeraer , 



Ichthyotaenia torulosa 



losD . 

T V\ "f" TTrt "f - «1 43 "l d 4~ #*> "K* il l rN o Q 

XOfl l> li,/0 wd-clllii bUi UlObd 

Zschokke , 


T /"» \\ *t~ "in TT/~\ "f~ O O V*. "1 n "f" ^\ 1 "1 ~1 /"\ O Q 

icii Oiiyo oaenia toruiood 

v. Ratz, 



Ichthyo taenia torulosa 

tfric & Vavr}, 1901:111-112 in 


Ichthyotaenia torulosa 

Qui. Schneider 1902:24 & 1905: 



Ichthyotaenia torulosa 

-L,uhe t 



SPECIFIC DIAGNOSIS. - Characters of the genus. Large 
cestcdes from 65-600 mm. long by 1.2-2.25 mm. broad. Head large, 
prominent, swollen, flattened dorscvent rally , without rostellum, 
without fifth sucker. Breadth of same 0.480-0.600 mm, f thickness 
about 0.300 mm. Suckers circular, prominent, with deep cavities, 
directed a^teriod and out. Diameters of same 0.18-0.20 mm. Neck 
thick, broad, 2-3 mm. Ion?, wrinkled. 

]Airst proglottides much broader than long, mature a.nd ripe 
proglottids almost always broader than long. Max. length of ripe 
proglottids l.Omrn^ max. breadth of same 2.5 mm. Proglottids 
fleshy, well delimited by deep inter-segmental furrows, corners 
of segments prominent. End proglottid rounded posteriorly. In 
its posterior end a deep indentation. 

GENITAL ORGANS.- Genital pore irregularly alternating, 
situated near middle of lateral proglottid margin. Vaginal opening 
dorsal and anterior to opening of cirrus-pouch. Cirrus-pouch and 
vagina opening into a common genital sinus. 

Male orga ns .- Testes large, 0.16x0.08 mm, 100-110 in 
number. Testes in two layers, the one occupying the whole dorsal 
field between vitellaria, completely covering ovary. Vas deferens 
all excentric mass of coils reaching from cirrus-pouch tc middle of 
proglottid. Cirru s-pcuch about 0.255 mm. long by 0.085 mm. "broad, 
extending l/4-l/6 across proglottid breadth. Cirrus short, not 
heav: . Ductus e jaculatoriu s in few or no coils. 

Fe-nale organs .- Typical of genus. Vagiral . openirg dorsal 
and anterior to cirrus-pouch. Sphincter vaginae very weak, near 


vaginal opening. Vagina not crossing cirrus-pcuch. Vitellaria 
voluminaus, follicles large and closely packed together. Ovary 
bilobed , lobes long, thick, and irregular in outline. Ootype and 
©Ocapt present. Organs of interovarial space as in genus. Uterus 
in ripe proglottids with 3-4 lateral pouches on either sice. Uter- 
ine pores not observed. 

EGGS.- Outer shell, hyaline, 0.055 iron, grarular second 
shell 0.032 mm. . embryo about 0.021 mm. in diameter. 


HABITAT.- In intestine 



: Locality : 

: Collector 

: Authority 

vy^^Ji x c v coco , 

i iJCI 1 ill i 

Bl ach ' 

1 Riiriolrhi (1819*1^0) 

Cyprinus Jeses : 

Greifs - : 

Bud ol phi : 

:Rudolphi (1819:150) 

Cyprinus orfusrldus mel- 
snctus orfus 

wald ! 

:Proelich : 

iRudolphi (1819:150) 

Jjuciscus leuciscus : 

Zeder : 

:E'Udoiphi (If 19; 150) 


Leuciscus leuciscus : 

v. Linstow (1878:254) 

•Leuciscus leuciscus : 


Fri¥ and : 

Friv & Vavra (1901 J 111- 

Leuciscus leuciscus : 

Bohemia : 
Lake : 

Vavra J 

Nufer : 

: 112) 
Uufer (1905:75) 

Leuciscus idus : 

'Lucerne : 

Abild- : 

Liesing (1850:559) 

Leuciscus idus ! 

M. C.Y. ! 

gaard : 

Diesing (1850:514) 

Leuciscus idus : 

Finland ■ : 

Gui. : 

Gui .Schneider (1902:24 

Leuciscus grislagine : 

Tullberg : 

& 1905:24-25) 
Lonnberg (1889:15) 

Alburnus bipunctatus ! 

Liesing (1850:514) 

Alburnus : 


iLiesing (1850:514) 

Aiuui nup lutxuuo < 

v T.instow (1878*258) 

Alburnus lucidus : 

Lake : 

Zschokke : 

[Zschokke (1884:20) 

Alburnus lucidus 

Geneva : 

:Lake : 


Kraemer (1892:55-71) 

Alburnus lucidus : 

: Lucerne 


•v.Linstow (1891:565) 

Alburnus lucidus : 


: Zschokke (1896:775) 

Alburnus lucidus 

: Lake 

: Nuf/er 

: Nufer (1905:75) 

Aspius rapax 

: Lucerne 

: Creplin 

iDiesing (1850:514) 

Ah ram. is brama. 

[V.Linstow (1878:258) 

Abramis brama 

: Lake 

• f 
v.Rat z 

v.Batz (1897:159) 

iQus meiaiioxus < 

: Balaton 

'V Linqtovw (1878*255) 

Pelecus cultratus 

: Lake 

: v. Eatz 

! V, Ratz (1897: 159 ) 

Gobis fluviatilis 

: Balaton 
: Lake 

: Nuf er 

: Nuiei \±y\JD. /d) 


: Lucerne 


Blicca bjoerk^a ' 


' Nufer : 

'JTufer (1905:75) 

Lucerne j 

Coregcnus fera 

:Lake : 

:Zschokke : 

Zschokke (1884:20) 


Coregcnus exiguua albellus 

: Lake 

:Nufer : 

:lTufer (1905:75) 

: Lucerne : 

Coregonus gchirgii helve- : 


:TTufer : 

Nufer (1905:75) 

t icus : 

: Lucerne 

Saline salvelinus : 


:Uufer : 

:Nufer (1905:75) 


Perca fluviatilis : 

Lake : 

Nufer : 

ITufer (1905: 75) 

: Lucerne 

Pri-f. & : 

Perca, fluviatilis : 

Podie- : 

Fri-f- & Vavgra (1901: 

Ttuead Bo- 

:Vavra : 

: 111-112) 

hemia J 

Lota vulgaris : 

Lake : 

' Zschokke 

Zschokke (1884:20) 

: Geneva 


HISTORICAL SUMMAEY/- Rudolphi (1810:111) gave a diagnosis 

and ym 

svnon and description of this form which is here quoted verbatim." 

Taenia tcrulosa Batsch. 
Taenia: capite t runcato , oscul is o rbic ula ribus marginati s, 
collo mediocri , articuli s crassiusculis ( subquadrat is ) subrotundi s . 

Bloch Abh. p. 11. Tab. 2. fig. 1-4. Taenia articulis rotun- 


Batsch Bandw.p.181 n.27. fig. 105-108. Taenia torulosa. 
Schrank Verz. p. 49. n.150. T. orbicularis. 
Gmel .Syst.n.p. 3081 .n.85. T. torulosa. 

Frolich im ffaturf. 25. p. 88 - 61. T. 3. fig. 4-6 T. simplex. 

Zeder Nachtrag p. 220. Rhytelminthus Cyprini . 

Rudolphi in Wied. Arch. 111. 1. p. 110. Taenia torulosa. 

Zeder Naturg. p. 352. n.39. Halysis torulosa. 

Hab. in ventriculc vel intestinis Cyprincrum. In Cyprin o 
Iese Bloch et ego Aprili, in Orf a Frolichius ecdem, m Leucisco 
Zederus Ma jo mense, reperimus. 

Descr. Vermes Zederiani septem ad novem lineas longi, 
dimidiam lati; Pr'olichiani duos ad quinque pollices longi, postice 
lineam lati; mei pedem non superantes, vix lineam dimidiam lati; 
Blochiani duos pedes lcngi, lineam et quod excurrit lati. 

Caput truncatum, depressum, sub motu pclymorphum, infla- 
tum et csculorum directionem sum mopere mutans; haec circularia, 
concava, marginata, margine simplici vel duplici, interdum occultato; 
mox omnia quatuor, mox due tantum in conspectum veniunt , ceteruin 


oculis nudis usurp an da. Collum depressum, mediocre, a capite 
crassiore discretum. Corpus crassiusculum , subaequale, margine 
crenatum, articulis confluentibus, margine pcstico vix incumbent ibus. 

Obs. 1. Blochius specimina sex maxima vidit, eorumque ar- 
ticulcs quam in meis magis discretes, ovis farctos, et foramina 
marginalia ( tamen ncn ulterius exposita, neque delineata) ob- 
servavit. Frolichiani vermes a meis vix ulla nota differunt, nisi 
quod collum longum dicat , quod ipse mediocre, Blochius breve voca- 
verit, hoc autem vermis totius raticne habita facile explicatur; in 

jvermibus enim bipedalibus collum vix pollicare Blochio breve visum, 
quod Frolichic in bivel quinquepollicaribu s longum, mihi in pedal- 
ibus mediocre fuerit. 

Obs. 2. Zederiani vermes valde pusilli fuere, ideoque a 

, nostris magis distant; ab osculis quatuor suctoriis totidem canales 
oriebantur, in ccrpore progressi, tandem in unicum vas abeuntes, 

I quod in cauda obtusc-acuta t erminabatur . Haec vasa mihi non visa, 
sed vermes illi tenelli plani, ideoque pellucidi fuere, nostri muto 

, crassiores, fere t eretiusculi . Zederus etiam postea ipse Bhyt elminthum 
cyprini clim sibi dictum ad T. torulosum pertinere suspicatus est, 

j quod clim in diario Wiedemanniano aeque indicaveram. Conf. etiiam Tae- 

| niam Cyprini Idi n.114. dictam. 

Rudclphi (1819:150) stated that this species had been 
found by Bloch, Proelich, Zeder and himself in certain species of 
the Cyprinidae. 

Dujardin (1845:584) did not record that he found this 
species. He collected his data from descriptions by Zeder, ?roelich, 


and Bloch. His description reads--" — Long de 16 a 20 mm, el large 

de 1 mm, 12 (Zeder) , ou large de 50 a 135 mm, et large de 2 mm, 25 

(Froelich), ou long de 330 mm, large de 1 mm, 2 (Bud), ou enfin 

long de 660 mm, large de 2 mm, 25 a (?) (Bloch); — tete trouquee, 

de forme t res-variabl e , que ses ventouses dcut le bord est 

saillant; trompe nulle; -- ecu de longueur mediocre; — articles 

assez epais, presque ronds." 

Diesing (1850:514) found this species in Leuciscus idus 

and in /lburrms b ipvnetatus and further stated that Creplin found it 


in A spin s rapax and in Alcurnus .piesing's diagnosis added a little 

to the previous knowledge of the form.* Taenia torulcsa Batsch, " 
" Caput l atuim depressum truncatum, acetabulis lateralibus limbo 
prominulo. Ccllum longum. Articuli t erst iuisculi subaequales longi 
confluentes. Aperturae genitalium. . Longit. 7-10"; latit. 4/z- 
l"'- . " 

Von Linstcw (1878 and 1889) catalogued the hosts from 
which this species had been collected. 

Zschokke (1884:20) stated that he found Taeni a torulosa 
Bud. in Coregonus f era ., Lota vulgari s and Alburnus lucidus from 
Lake Geneva. Since Zschekke reported Taenia lengi chilis and T. 
ocellata from Coregonus f era in addition to Taenia torulosa it is 
highly probable that bis specimens belonged to one of these species. 
Zschckke furthermore states that his specimens were too young to 
permit him to recognize ary trace of internal organs. These state- 
ments apply equally well to his parasite report on Lota vulgaris 
in which case he found Taenia ccellata in addition to T aenia tornl o- 
sa . 


His feu** young specimens from A lburnu s lugidus may have been 
Taenia^orulosa if only the question of host be considered. Of 

Is. s / 

t^es'e four specimens he wrote: "La tete etait large, tronquee, 

v / 
les ventouses tres - fortes et saillantes. Le cou etait de longeur 

medicare, la segmentation en articles pea. accusee." 

Lonnberg (1889:15) reported the finding of Taenia torulosa 

by Professor Tullberg. He gave nc description and did not state the 
locality. Vcn Linstow (1891:565) found Taenia torulosa in Alburnus 
lucidus . TT e rave no data. 

Kraemer (1892:55) found Taenia torulosa in Alburnus lucidus 

Lake Lucerne. Kraemer examined more than 150 specimens of Coregon m 

fera without finding a single specimen of Taenia torulosa and 

among numerous specimens of alburnu s lucidus but a single host was 

infected with Taenia torulosa Kraemer made the first careful 

morphological and histological study of this species. Lonnberg 
/ ~ ~ x . m torulosa 

(1894:801-3) included Taenia ~T in the llst °f species in 

his new genus I chthyo taenia . 

Zschokke (1396:775) found Ichthyotaenia torulosa (Bat sch) 
in the intestine of LeuCc i scus leuQpiscus L. He found it in no 
other hosts. 

V. Ratz (1397:159) reported finding Ichthyotaeni a t orulosa 
in Abram-i s braag L. and in Pel ecus c ult rat u s L- in Lake Balaton, 
Hungary., but he gave no description. 

"Fric and Vavra (1901:111-11^) reported what they considered 

to be IcbthypLtaenia t o rulosa (Batsch) (misspelling for torulosa) 


from Leuciscus l euciscus end Perca f luviat ilis Podiebrad , Bohemia. 



The description their specimens agrees very well with that of 

Prot eocephalus percae and a part or all of their specimens may 

belong to that species. The Hasling ( Leuciscus leuci scus ) would 

be a new host for P. percae and it may be that their specimens from 

that host are as they identified them. P. t brulcsus . Moreover, it 

is not . that FriT and Vavra failed to distinguish between the two 
species of cestodes^P. percae and P .tor utiosus which were present in 
Perca fluv iatilis and Leuciscus leuciscus respectively. It is 
impossible to make a complete determination from their data. 

Gui. Schneider (1902:24) found Ichthyot aenia torulosa in 
Leuciscus idus in Finland. Three years later he (1905:24-25) 
briefly but concisely described this species froro. Leuci scus idus . 
His description agrees very well with that of Kraemer (1892). 

Fufer (1905:75) reported Prot eccephalu s torulosu s (Batsch) 
from Perca fluviatilis , Alburnus lucidus^ Squalius leuciscu s, 
Elicca bjoerkna , Gobio fluviatilis , Coregcnus exiguus albellus , C . 
scMnzi i helveticu s, Salm o salvelxinu s. Some of IsTufer's report is 
open to doubt. In 59 specimens of Perca fluv iat ilis not only he 
failed to find the species which Zschokke found in that host but he 
found P. longicqllis and P.^ orul&sus , a species which only once 
before (Fric and Vavra 1901) had been reported from that host." In 
that instance I called attention to the fact that the specimens in 
question were probably P. percae not P. torulosus . Nufer's P. 
to rul o su s and his P. lcngi chilis may have been P. percae . 

> T ufer f s next four hosts after Perca fluviatilis a re all 
Cypri^idae, from which group hitherto this species has been report- 
ed. It is extremely doubtful if the Salmonidae harbor this 


species. Zschokke who in 1884 reported young specimens of Taen ia 
torulcsa from Qoreg-onu s fer a, in 1896 found this parasite only in 
Alhurnus lucidus . Zschokke 1 s paper (1896) contained the results 
of his investigations on more than 1600 fish from Lake Geneva and 
the ffhine^hence his data was fairly comprehensive. Moreover, from 
the three species of Salmonidae in which he claimed to have found 
P. t orulosus "Fufer reported two other species of Prot eocephalus ^ 
P. longic % l li s and P. ocell atus. Here again Nufer probably made a 
misdetermination. I have shown that Fufer's work (see discussion 
of P. macrocephalusfpage ) to be untrustworthy anct^the de- 

termination of this species there seerns to be no exception. That 

7T ufer made a midetermination of his P. longicjjllis is not at all 
improbable since specimens labelled Taenia longicallis aus Porelle. 
which have been received by Professor Ward from Professor Zschokke 
have proved to belong to a different species. ITufer's tabulated 

description of P. longicallis (ITufer 1905:147) does not agree in 

important points with v . Linstow's description yet it is very 
manifest that he drew on v. Linstow's description for a large part 
of his data. Likewise parts of ITufer's description of P. t orulosu s 
(JTufer 1905:147) fail to agree with the descriptions of that 
species by Schneider and Kraemer. 

Luke (1909:32) gave a short specific description of this 
form under the name of Ichthyetenia torulosa (Batsch) , 

MATERIAL.- The following study is based upon some of fi. 
Schneider's specimens which Prof. H. B. Ward secured from Professor 
Levander at Helsingfor&. This material is labelled 9 1 chthy o taenia 



t orulosa Batsc^. 

leuciscu s idu s. Porkala. Juni 1901. det . G. Schneider." 
It now bears the number 10.121 in Professor Ward's collection. 
Prom this material frontal and transverse sections have been made. 
Some heads have been studied in glycerine. 

A careful comparison of Schneider's (1905) and Kraemer's 
(1692) descriptions shows very gccd agreement. Zschokke's (1864) 
description in parts agrees pretty well witBa. these two. Lufee (1909] 
seems to have based his specific description largely on the work 
of Zschokke, Kraemer, and Schneider. Data from these sourses is 
used in this description. 

EXTERNAL CHARACTERS. - This is one of the larger species 
of Proteoxiephalus infesting fish. In length it varies from 65-600 
mm. Specimens of the latter length are rare and have only been 
repor T ed by some of the earlier investigators. The longest com- 
plete strobila examined by me was 110 mm long. The breadth of the 
strflbila var ies from 1.2-2.25 mm. 

The head (PI. ^ fig. 7^ ) is large and very prominent, 
somewhat swollen iy\ appearance. It is flattened dorso-vent rally , 
and is somewhat flattened at the apex.- Schneider found it 0.500 
mm. broad. My measurement of the head was 0.450-0.600 mm. broad 
by about 0.320 mm. thick. The suckere are very prominent, nearly 
pi<eullar_ in outline, with deep cavities and strong musculature. 

They are directed anteriad and a little out. The suckers are 0.200 
mm in diameter according to Schneider and about 0.180 » •Q.J.OO mm 
in the heads examined by me. A fifth Bucker is not present. In 


sections of the head I was unable to find even a trace of an end- 
organ. Nufer (190^ ;.147) stated that the head of a specimen 37 

mm^ measured 0.255 mm. broad and the suckers of the same were 0.10 
mm in diameter. 

T^ufer's measurements of the head do not agree at all with 
Schneider's or my own. Neither Kraeitier nor Zschokke give measure- 
ments of the head and suckers hut they state that the head is 
large, the suckers large and round. Nufer's specimens which come 

from Perca f luviat il i s most certainly could not have been P. 

f all ftx 

t o rul o su s .oatscb but were either P. percae . P, r - ,9 n, sp. or 

an undescribed species. 

The neck which is thick is about 2-3 mm. long by about 
0.20-0.30 mm. brocd. It passes over gradually into the first 
proglottids which are much broader than long. Mature and ripe 
proglottids are almost always broader than long. The max. length 
observed was 1.0 mm and the max. breadth 2.5 mm. Many nearly ripe 
proglottids measure about 0.29 mm. long by 1.30 mm broad. The 
proglottids are thick and fleshy with well defined limits. The 
indentations between the segments are deep. The corners of the 
proglottids are quite pronounced. The end-proglottid is rounded 
posteriorly. In its posterior end it has^deep indentation into 
which the excretory ducts empty. 

EXCRETORY SYSTEM . - According to Kraemer the excretory 
ducts in the head and neck region are very prominent with many 
anastomosing coils. In the neck he found many branches of the 
excretory vessels leading to the exterior. My sections failed to 


show these relations. 

GENITAL ORGANS.- The genital openings are irregularly 
alternating, and are situated one in or near the middle of the 
margin o^ each segment. The vagina opens not strictly anterior to 
the cirrus-pouch hut somewhat dorsal thereto, cirrus-pouch and 
vagina open into a common genital atr^^m. Kraemer states that the 
vpp-,ir\8 and cirrus-pouch open near each other but not into an 
at rium. 

Male Organs . - The testes are large, spheroidal or by 
pressure sometimes rendered polyhedral. They measure / according to 
Schneider, about 0.160 mm. in sagittal by 0.080 mm. in frontal di- 
ameter. My measurements are about the same as these. The testes 
lie in two layers. The dorsal layer (PI. V fig.'7>~* ) covering 
the entire area founded by the vitellaria and the anterior and 
posterior margins of the p(S^/glott id. The ovary is covered complete 
ly. In the dorsal layer there are about 70 testes. In the more 
ventral layer there are about 30-40 testes. Hence the total number 
of testes is about 100-110. Kraemer' s drawing (PI. H fig. 77 ) 
shows quite a small number of testes but shows the ventral area 
of the worn and so cannot show but a small number. 

The coils of the vas deferens form a thick mass extending 
from the cirrus-pouch tc the mid-field of the segment. Kraemer 
figi res the coils of vas deferens lyin^ in the middle of the 
segment . 

The cirrus-pouch is short and relatively thick. Its 
length is about 0.255 mm. and its breadth 0.085 mm. The ratio 01 


of its length to the proglottid breadth is about 1*4 according to 
G. Schneider, l/3 in Kraenier's drawing, and l/4-ljfe in my prepara- 

The cirrus- is short and not very thick. Kraemer's 
description of the cirrus is faulty in that he said that it 
was armed with hocks which are recurved and extend back into the 
tissue of the cirrus itself. There are no bocks. I bave not seen 
the cirrus protruded. 

The ductus e jaculat orius forms but a very few ceils or 
instead of ceils it may lie in sinuous curves within the cirrus 

Female organs .- In their general relationships these are 
typical of the genus. 

The vaginai which opens mostly dorsal tc the cirrus-pouch 

bos a very weak sphincter muscle situated very near the opening into 

the atrium. This sphincter vaginae is made up of a few strands of 

circular muscle fibers which are easily overlooked. In its course 

to the int er-ovariel space the vagina dees not cross the cirrus- 

smo.1 1 

pouch but passes dorsal and anteri^ad of the pouch. A. receptac- 
ulum seminis is present just anterior to the ovary. 


The vitellaria are later;!^ v olummcus , end follicular. 
The follicles are large and closely paccked together. The bilobed 
ovary is large, well developed, thick and irregular in outline. 
The lobes are net as slender as they are shown in Kraemer's figure. 
In prcrlottids 1.3 mm. bread the cvary may have a span of 0.80 mm. 


and the lobes may be 0.130-0.140 mm. thick. 

A Muscular oocapt and an ootvpe are present. The organs 
of the in> erovarial space are arranged as in other species cf the 
genus. The uterus in ripe proglottids has 3-4 lateral out-pocketing 
on either side. The uterine pores have not been observed. 

EGGS.- These were not described by Kraemer and Schneider. 

The outer shell is thin and hyaline. It's diameter measures 0.055 

mm. The granular second shell has a diameter of abcut 0.032 mm. 
the - 

and^ embryo about 0.021 ram. a delicate membraree invests the 

R elationship s 

P. torulosis s differs from, many other species of the genus 

by its large size, and from a large number of the species thru its 

lack cf a fifth sucker. In max. length and breadth of its strobila 

it is the largest species cf Prot eocephalus yet described* In the 

width of the head and diameter cf the suckers it is exceeded only 

by P. umbl cpliti s and P. perplexus . Yet in observed length P. 
greatly exceeds these species. P. tomlosus. 

t c nil o su s ^ may be difgrent iated from these species by its lesser 

number of uterine pouches, its weaker sphincter vaginae, its shorter 
cirrus-pouch, its double layer of testes a.nd in the arrangement cf 
dorsal layer of the same. It is greatly different from the other 
TT. American forms thus far described. Among the IDuropean forms 
P. ^?or u.lo sus i s approached in size only by P. longicola is . _P. 
longlc e ll i s # however .has a well developed fifth sucker and, according 
to von Linstow^a much smaller mumber of testes. P. t orulosus may 
be differentiated from P. perca e and P. cern uae by its larger head, 
la-ger suckers, l-ack of a f if th 4icker, more numerous testes and 


the different arrangement of the same, and by its smaller number 
of uterine pouches. P. t or ul osus is much larger than P. macrcce ph 
alus . It has a larger head, larger suckers, fewer uterine pouche 
and a different arrangement of the dorsal layer of testes. 

P. t orulosus is so different from the other described 
species of Prot eoceph a lus that a danger of its being confused with 
them is scarcely possible. 


(PI. / Figs.L/^ 
1906: I chthy'. taeni a pe^ t astor ttf^Klaptocz , 1906". 130-133. 

SPECIFIC DIAGNOSIS.- Characters of the gerus. Cestodes 
of small size. Strobila up to 28 mm, long; max. breadth of same 
1.19 mm. Neck none. First proglotcids many times "broader than 
long. Mid-proglott ids 1.19 mm. "broad 0.150 mm. long. Last 
proglottid 0.59 mm. broad^0.21 mm long. Head large, oval in cross- 
section, 0.975 mm. "broad x 0.83 mm. thick. No rostelluin, no hooks. 
Pour large suckers, oval or round in outline^ 0.50 mm. in diameter 
situated ir. four quadra,nts. Suckers separated from each other by 
longitudinal furrows. A true fifth sucker 0.085-0.100 mm. in 
diameter at apex of head. 

GENITAL ORGANS.- Genital aperture about middle of 
proglottid. Vagina posterior to the cirrus-pcuch. Testes 70-100, 
arranged irregularly between vitellaria vvith tendency tovvard two 
fields. No data on cirrvs-pcuch and vas deferens. Vitellaria 
lateral, not voluminous . Ovary bilobed, posterior. Uterus immature 

HABITAT.- Pclypt erus bichir Geoff r, White Nile near Khor 


MATERIAL.- A single specimen was taken by Dr. P. Werner 
Peb. 16, 1905 while making a journey up the Nile river into the 
Sudan . 

DESCRIPTION .- All oata are taken from Klaptocz (1906) 




the wo rm are 



Length of complete strobila 

28.0 mm. 

Later&l diameter of scolex 



Dorsoventr&l diajneter of scolex 



Suckers, max. dia.vrieter 



Fifth sucker, diameter 



Proglottids near scolex, 







1 ength 



A proglottid near the middle cf strobila, 

max. breadth 






ProglottiQ, the sixth from the end, 

oi eadth 






Proglottid, next to last, 


. 410 


1 ength 



Last proglottid, 


. 330 






(/>/•//* ^ 

The scolex^is somewhat oval in trans-section, surpassing 
the breadth of the first proglottides. It possesses no rostellum, 
nc armature of hooks but has an apical sucker 0.085-0.100 mm. in 
diameter. The four equal sized suckers of somewhat oval or round 
shape, of varying size, about 0.50 mm. in diameter, are situated on 
the four quadrants of the head. The four quadrants are somewhat 
swollen and are separated from each other by longitudinal furrows. 

The apical depression is a true sucker, as shown in the 
stained specimen. 

There is no neck, the first segments appearing under the 
microscope 0.060 mm. behind the sucker margins. 

The proportions of the proglottids are best given in the 
table of measurements. From the drawing the st rooilat ion is evident 
the posterior angles of the proglottids being quite prominent. 

GEMITAL OKGAFS.- The anlagen of the sex organs may 

be seen 7 mm. back of the scolex-tip. The testes are well developed 
10mm. back of the scoleix* j . These are elliptical in shape, their 
long axis lying in the long axis of the worm. They number 70-100, 
lie irregularly scattered between the vitellaria and anterior to the 
ovaries. They are most numerous in the lateral fields but they also 
occur in the median part especially in the anterior region. 

Fo data is given concerning the cirrus-pouch and the vas 


The genital pore is irregularly alternating. It is 
situated near the middle of the margin of the segment, but in the 
last proglottids a trifle anterior. 


The vagina opens posterior to the cirrus-pouch. Just 
"before its opening it possesses a considerable broadening, the 
diameter of which is about equal to the "broadest diameter of the 
cirrus-pcuch. On either side of this broadened part the vagina is 

The vitellaria are lateral longitudinal glands which 
develop late. They are not voluminous. 

The bi-lobed ovary lies in the posterior part of the 
proglottid as in other species of the genus. 

The uterus was immature in his specimen. This species 
easily separates itself from the most of the forms from fish by 
the large size of its head and suckers. The position of the vagina 
posterior to the cirrus-pouch is a further diagnostic character of 
value. The lack of an unsegmented neck and the proportions of the 
prcglottids are further characters which serve for its identification 

Unfortunately Klaptocz failed to describe and to figure 
the cirrus-pouch, cirrus, and vas deferens which are of great value 
in making a positive determination. The uterus being immature 
could not be described. For these reasons this species which is 
very likely a species of P rpt eocephe lus cannot be accurately placed 
in its relations to the other members of the genus. 



(pi.f iigB,!f7-/*rJ 

1896: Ichthyotaenia fossata Riggenbach, 1896. 166-193. 

SPECIFIC DIAGNOSIS.- The characters of the genus. 
Cestodes of short length, 5.5-4 cms long. Scolex large, visible to 
naked eye, max. breadth of seme 0.714 mm. Anterior end of scolex 
conical, bearing at apex a slight concavity, not a fifth sucker. 
Suckers, round, 0.34 mm. in diameter, home at broadest part of head. 
Neck broad, 0.85 mm. long. First prcglovtt ids broader than long, 
0.612 mm. broad x 0.155 mm. long. Mature and ripe proglottids, 
quadrate to longer than bread. End proglottid not observed. Lateral 
margins of proglottids quite straight. About the genital sinus a 
slight elevation or genital papilla. 

Genital organs .- Genital aperture marginal, irregularly 

alternating, situated a little anterior to the middle of the 

proglottid. Testes, numerous, 120-lSO^ in medullary parenchyma 

between vitellaris. Vas deferens, a loose mass of coils between 
and middle of proglottid, posterior to cirrus-pouch 
cirrus-pouchi Ductus ejaculations with few coils in cirrus-pouch. 

Cirrus short and thick. Cirrus pouch pear-shaped, 0.30 mm. long, 

extending about l/3 across the proglottid breadth. 

Vagina, opening anterior to cirrus-pouch. Vaginal sphincter 

small, near opening. Lunen of first part of Vcgina broad, nearly 

as large as cirrus-pouch. No coils in vagina anterior to ovary. 

Arrangement of organs in interofarial space typical of genus. Ovary 

bilobed, posterior, lobes connected by midpiece. Vitelline glands 


follicular, lateral, extending full length of segment. Uterus in 
ripe proglottid with many lateral outpocket ings on either side. 

EGGS; round thin shelled structures, size not given. 

"HABITAT.- Intestine of Pime lodus pati valine i Rio 
Paraguay, S. America. 

The material was collected "by Dr. Ternetz in January and 
February 1894. The following description is hased on Riggenbach' s 
(1896) paper. 

EXTERNAL CHARACTERS.- The worms as collected were of 
small size, 3.5-4.0 cms. long. There was no undamaged strobila. 

'The neck is 0.85 mm. long and is quite broad. It passes 
imperceptibly into the youngest proglottids. 

The scolex^is relatively large, and is perceptible to the 

naked eye. It bears four large suckers at its broadest zone. Here 

the scolex^ 0.714 mm. bread. Prom this point the scolex becomes 

smaller toward the^end and also toward the neck. Its anterior 
region is conical. At the apex of the head is a slight depression 
which structurally is not a fifth sucker. There is r.o rostelT2uni. 

The suckers are round in outline and they measure 0.34 mm. 
in diameter. 

The youngest proglottids in the proliferation zone are 
narrow tranverse bands 0.135 mm. long by 0.612 mm. broad. With 
increasing age the proglottids elongate until mature and ripe 
proglottids may be quadrate and the oldest proglottids even longer 
than broad. An end proglottid was not seen. 


The margins of the segments are nearly straight. At the 

place of the opening of the genital passages the surface is slightly 
raised to form a small genital- papilla. 

I5TTEK6TAL ANATOMY. - Excretory System.- In the head the 

excretory vessels run close together. Wear the suckers a circular 

E anastomosis join? the vessels together. In the space between the 

suckers is a complicated plexus . 

The two pairs of main lateral excretory vessels run thru 

the length of the worm in^nearly straight course. In the posterior 
margin of each proglottid is a transverse anastorasis connecting the 
main vessels. In this region also and in the neck short ducts arise 
from the main ventral vessels only and pass to the exterior, opening 
on the surface near the posterior angle of the proglottid. The 
lumen of such ducts becomes smaller near the periphery of the worm. 
The passage thru the cuticula is quite small. 

GENITAL ORGANS.- The genital aperture is marginal, 

i rregularly alternating, situated a little anterior to the middle 
of" the proglottid. A very small genital papilla is present. This 
is due largely to a thickening of the margin of the sinus. 

The anlagen of the genital organs appear in about the 
twentieth segment. , 

Male organs ." The testes^are numerous, 120-150, and re- 
latively small, 0.5 mm. in diameter. They are round or polygonal in 
outline. They fill the whole field between the two vilelline glands 
and the anterior and posterior margins of the proglottid. 

The vas deferens forms a loose -lass of coils between the 


cirrus-pcuch and the middle of the proglottid. The mass lies 
posteriad to the cirrus-pouch and is excentric lying on the 
porase side of the proglottid. 

In the cirrus-pouch the ductus e jaculatorius forms a few 
coils and then it passes over into the cirrus. The cirrus is en- 
larged into a thick vesicle which fills up nearly the outer half 
of the cirrus-pouch. 

The cirrus-pouch, which is about 0.30 mm. long ; reaches 
about l/5 across the proglottid breadth. It is an elongated oval in 
shape . 

Fesiale organ s- The vagina which always opens anterior to 
the cirrus-pcuch has a small sphincter near its opening. Further 
within, the lumen of the vagina enlarges into a cvoidal vesicle 
which may be almost as voluminous and as long as the cirrus-pouch. 
The vagina forms no coils anterior to the ovary. Riggenbach found 
no receptaculum seminis but he found a widening of the vagina 
anterior to the ovary. The two dilatations of the vagira take the 
place of the reeeptaculum seminis. 

The arrangement of the organs in the int e^ovarial space 
is typical of the genus. In this space are located the ocCtype, 06- 
cap^shell- gland , cviduct, coils of the vagina, vitelline duct, and 
the uterine passage. 

The ovary is a bi-lcbed structure, situated in the posterio 
part of the proglottid. 1 The ovary does not seem to be made up of 
blind- pouches, as is true in the ovaries of most of the Taenias. 
The outline is somwwhat irregular, due to the sac--like processes 



which may be st: i on the plump mass of the lo e. These extend 
poster! ad nearly to the proglottid margin end laterally to the 
excretory vessels. 

The vitellaria as in all Prct eo eepha.lids are made up of a 

large number of single follicles which are arranged in long hands 

the the 
in each lateral field of, proglottid. They extend full, length of 

r A 

the proglottid. The vitelline ducts arise in the posterior region 
of the segment and their course to the inter-ovarial space is like 
that of other Prct eccephalids . 

The shell-gland is poorly developed. The uterus lies in 
the middle of the proglottid. "As a canal with numerous lateral 
branches it extends itself throughout the whole proglottid." 

EGGS.- "The uterine eggs are round thin shelled structures." 
Size not given. 


PB0T1OCBPE&LUS ESOCIS (Gui Schneider) 
(Pl.^Eigs.^ $~3l). 
1905: iCchth yota enia eeo cia Gui. Schneider, 1905. 

SPECIFIC DIAGNOSIS.- The characters of the genus. Cestcdes 
small and slender, 1-4.7 cms long, max. "breadth 0.7 mm. Scolex 
0.140 mm. bread, 0.105 mm. thick. No fifth sucker, no end organ. 

Suckers shallow, weak, small, abcut 0.050 mm. in diameter, 
directed anteriad. Diameter of sucker opening 0. 026-0. 03k; mm. Feck 
2-5 mm. long. Number of proglottids in specimen 4.7 cms long 66 
youngest proglottids twice as broad as long 0.370 mm. Broad and 
0.170 mm. long. Middle proglottids broader than long, 0.680 mm. 
"broad x 0.390 mm. long. Last proglottid longest, 0.650 mm. long x 
0.460 mm. broad at anterior end. 

Segmentation not evident. Transverse intersegmental 
furrows shallow. Surface smooth. 

as in genus 

Genital organ s: - Genital pore^not cn genital papilla. 
Pore marginal, irregularly alternating, about middle of proglottid. 

Fema le organs: - Vagina anterior tc cirrus-pcuch. Near 
vaginal opening a small sphincter vaginae. Re eeptaculum seminis 
anterior to ovary. Passages in int erovarial space as in genus. 
Ovary bi-lobed, lobes ovoidal vitellcria sparse, follicles small. 
Uterusnot observed. 

Male or gans : - Cirrus-sheathf relatively long, extending 
nearly to middle of proglottid. Length of cirrus-pouch 0.230-0.250 
mm. Cirrus s^rnuous.- Vas deferens a mass of coils in middle of 


proglottid. Testes. 30-38 visible in mature proglctt ids , 44-52 in 
young prcglott ids. Testes arranged in tv c partial layers betv/een 
vitellaria. Dianeter of testes 0.050-0.090 mm. 

Excretor y vessel s : - Two pairs, ventral and dorsal. 
Dorsal vessels small, ventral vessels much, larger. 

HABITAT: - In intestine of Esox lucj qs L . Reval, Estland. 


SOURCE OF MATERIAL:- This species from Esox lucius .taken at 
Reval in Esthonia, Apr. 30, 1904 is briefly described by Gui. 
Schneider (1905-7). Thanks to Dr. H. B. Ward I have seme Schneider's 
preparations and alcoholics for examination. The paucity of alco- 
holic specimens prevented the use of the section method. I have 
therefore teen compelled to be content with a study of toto prep- 
arations and specimens cleared in glycerine. The description 
following is based largely on Schneider's data the I have^ wherever 
possible^ introduced my own findings. 

EXTERNAL CHARACTERS: - The worms are small and slender 
1-4.7 cms. long. I observed a maximum breadth of about 0.7 mm. 
17 o worms had ripe proglottids. 

The diameter of the scclex "is about 0.140 mm. The four 
shallow suckersvdirected partly forward, are weak and small and are 
about ^.05 mm. in diameter. An endcrgan is lacking^ tno perhaps a 
slight thickening of cells in the anterior part of the head is to 
be considered as the vestige of the shrunken end organ (Schneider). 
My measurements of a head and its suckers are: breadth 0.140 mm., 
thickness 0.105 mm, diameter of suckers 0.053 mm, diameter of 
sucker openings 0.026-0.032 mm. A most careful search of Schneider's 
toto preparation and of the specimen cleared in glycerine failed tc 
reveal a trace of a fifth sucker or of an endcrgan. The last might 
not have been visible under these conditions. A fifth sucker 
certainly is not present. The suckers are quite promirent. The 


sucker cavity is shallow. The 

slightly depressed head possesses a. short flattened tip. The head 
is not well set off from the neck which follows* 

Schneider says that the length of the neck is ah out l/3 
the length of the body. I find it to be from 2.5-3*4.5 mm. 

Schneider's largest specimen, 4.7 cms. long, was made up 
of 66 proglottids. The youngest proglottids were about twice as 
broad as long, the middle ones somewhat broader than long, and the 
posterior ones quadrate or a little longer than bread. The inter- 
segmental furrows were not plain. 

I found the measurements of the proglottids in a glycerine 
specimen to be as follows: first proglottids 0.370 mm. broad x 
i 0.170 mm long, middle and mature proglottids 0.680 mm. broad by 
0.390 mm. long. The last proglottid was the longest in the strobila 
0.650 mm. long x 0.460 mm. broad at anterior end. 

The segmentation is not evident. The proglottids are 

joined by their full breadth and the transverse furrows between 

the segments are very shallow. Ho longitudinal furrows,, present . 
of the 

Figures 3 three regions, show the proportions of the segments, 
anterior, pest-middle, and posterior end. (PI. Z igs. 3/*>,c. ) 

GENITAL 0RGA2TS: - The genital pore is not marked by a 
papilla. It is marginal, irregularly alternating in the strobila, 
and is situated about the middle of the proglottid. 

Male organs: - The cirrus- sheath is relatively very long 
and slender for it extends to the middle of proglottid or very 
near to the middle. Its length in the broadest proglottids is about 


0.230-0.250 mm. Schneider reports that its inner end lies very 
near the dorsal muscle layers. This is also the case in P. percae , 
P. pittgu&B P. exigH-tts and a number of other Prot e c c ephalu s species. 

The cirrus runs thru the cirrus-pcuch in a sinuous course, 
without the formation of a vesicula seminal is. The vas deferens 
forms a fairly large mass of ceils in the middle of the segment. 
This mass however is much smaller than in P. fallax Dj s p 
I e or in F. exigu qs, 

Schneider found about 30 testes in each segment lying in 
a single layer, about 8 in a transverse jhew and they measured 0.080- 
0.090 mm. in sagittal by 0.040-0.050 mm. in frontal diameter. 

I found it. difficult to count the testes in the more 
mature prcglottids.p/.^f ig.30 Shows 33 testes. In the younger proglot- 

tids vhere they are smaller and more easily distinguished I 
counted 44-48-52. The testes seemed to be in two partial layers 
filling area between vitellaria.. The larger testes measure up to 
about 0.070-0.080 mm. tho the larger number measure about 0.050 mm. 

Schneider found the diameter of the two ventral excretory 
vessels to be about 0.015 mm. and that of the two dorsal about 
0.0025 mm. 

Female organs : - The vagina opens anterior to the cirrus- 
pouch.Very near the opening, 0.030 mm. from it, is a sphincter 
muscle 0.015 mm. thick^according to Schneider. I was unable to 
observe this. From the opening the vagina passes as a tube with a 
wide lumen to the middle of the proglottid in a long smooth curve 
then it bends posteriad and passes back to the int epovarial space. 


As the vagina nears the middle of the segment the lumen becomes 
constricted. Early in its course it crosses the cirrus-pouch 
diagonally. The vagina is dorsal to the uterus. Schneider found a 
small receptaclum seminis just anterior tc the ovary. 

According to Schneider the passages In the int ei~ovarial 
space bear the relations customary to Prot eccephalids . 

The ovary is bi-lobed, the lobes being like two elongated 
balls connected by slender arched midpieces. 

The vitellaria are lateral and follicular. They are 
sparse. The individual f elides are small. 

The _nterus was not observed by Schneider. It is not 
well developed in any cf the specimens examined by me. Schneider 
saw the opening of the uterine passage and the preformed uterus 
opening just a little posterior to the middle of the proglottid. 

EGGS:- Not seen. Ho prcglottids were ripe. 

ReJ^aJU.£nsbjj)S.- This species is quite closely allied 
to the P. ambiguus (Dujardin) described by Gui. Schneider (l905-» .} 
However it is differentiated from P. ambig uus by its - much 

longer cirrus-pcuch and by its larger testes. P. ambiguus shews 
a slight genital prominence upon which the genital pore is 

It differs very markedly from the P. pingttj s found in our 

If. American Esox» lucj ne . The P. pingits is a lar ger worm with a 
— — _ — ______ » _ — | 

larger head, larger suckers, a functional fifth sucker, a relatively 
shorter cirrus-pcuch, a larger number of testes. 

P. fall ox , P. exi guu s , P. pusillus and P. percae are 


readily distinguished from P. esoci£ by their fifth sucker and by 
many other characteristics which are best shown in the key and the 
tables . 

P. macro c ephalus , a foim without a fifth sucker, is 
much larger, has more testes, a relatively shorter cirrus-pouch and 
in other ways also it differs widely from P. esocis. 


? 1319: Bothriocephalus barbatulae Rudolphi, 1819: 144- 
? 1850: Bothriocephalus barbatulae 
1872: Taenia 

1878: Taenia 

1891: Taenia 

1896: Ichthyotaenia 

1909: Ichthyotaenia 

Piesing, 1850:608. 
sagitta Grimm, 1872: 240-243. 

sagittat a Von Linstow, 1878:260. 
sagittata Monticelli, 1891:169. 

gittata Riggenbach, 1896:267. 
sagitta Luhe > 1909:33. 

SPECIFIC DIAGNOSIS.- Strobila small, up to 45 mm. long 
by 1 mm. broad. Head thick, ending in an unarmed point. Suckers 
four, in pairs on the basal part of the head. Suckers transversely 
elongated and heavily muscled. Heck up to 12 mm. long by 0.5 mm. 
broad. Proglottids few. Youngest proglottids about quadrate, 
older ones longer. Last eight proglottids 1.4-1.5 mm. long by 1 
mm. broad by 0.3 mm. thick. Last proglottid always rounded 

Cirrus-pouch large and clubshaped. Cirrus quite thick. 

Uterus with uterine pouches, number; , Genital-pore 
situated near middle of lateral margin of proglottid. 

HABITAT . - In intestine of Cobitis barbatula ^Petersburg , 
and Province of Novgorod. 

HISTORICAL SUMMARY. - Grimm (1872: 240-243) described 
some cestodes which he had collected from the intestines of Cobitis 
barbatula at Petersburg and in the province of Novgorod. These 
he named Taenia sagitta. He called attention to the fact that 


Rudolphi (1819:144) had proposed the name Bothicocephalus barbatula€ 
for seme cestodes from Cobit is barbatula. Rudolphis gave no 
diagnosis or description. Grimm also called attention to Diesing's 
(1850:608) reference to Bothriocephalus barbatulae and that Diesing 
gave no description of the form. 

V. Linstow (1878:260) listed Taenia sagittat a Grimm (for 
sagitt a) as a parasite of Cobit i s barbatula . Monticelli (1891:169) 
included Taenia s agitt at a (for sagitta) in a list of cestcde 

species parasitic in fish. Riggenbach (1896:267) included Ichthy- 

jet aenia sagitt at a (Grimm) in a list of I chthyo taeni a. Luhe- (1909: 

33) gave a very short diagnosis of Ichthyotaenia sagitta (Grimm) 

which he apparently derived from Grimm's (1872) paper. Because of 

its inaccessibility to many scientific workers^ Grimm* s (1872) 

entire description of this species is quoted. 

f/ TASEJTA SAGITTA.- Der beruhmte Helmietfholog Rudolphi ere. 

wahnt in seinem Werk Entozoorum Synopsis S. 144 eines Bandwurmes, 

den er im Darme von Cobitis barbatula aufgefunden hat. Da er aber 

nur ein Fragment, nicht aber ein voiles Examplar des Wurmes besass, 

so gab er auch keine eingehendere Beschreibung dieser als neu von 

ihm erkannten Art, sondern stellte sie graa$ zu den Grubenkopfen 

und belegte sie mit dem Namen Bothriocephalus barbatulae. Diesing 

stellte diese Art in seinem Systems helminthum zu den species dub- 

iae und setzte fu dem von Rudolphi gesagten nur noch hinzu, dass 


"Fragment a acephala servantur im M. C. V." (Syst. helm. I. p. 608). 
Etwas umstandlicher spricht sich S. Leuckart aus uber das Aussehen 
dieses Felminthen, indem er sagt , "an dem sehr kleinen kopflosen 


Stucke, das ich sah, war nicht s Ausgezeichnetes. Die Glieder langee 
als breit, Oder vollko-nmen quadrat isch. Keine ecUtlichen Ovarien" 
(Sig. LBuckart, Zoologische Bruchstucke. I. p. 57). Das ist Alles, 
was wir von diesem Bandwurme kennen und deshalb wird wohl nicht 
uberflussig sein, wenn ich im folgenden die Resultate meiner anatom- 
ischen Untersuchung kurz zusammen fassen werde, die ich an diesem 
Wur.ii unlangst angestellt habe. 

Ich fand namlich hier, in Petersburg, und im "!cwgorod r - 
schen Gouvernement 3 erwachsene Eaemplare des Wurms in Cobitis 
barbatula, nachem ich einige Hunderte dieses so haufig in unseren 
kleineren Fliissen vorkommenden Fischchens durchmust ert hatte. Aus 
dem Gesagten geht schon hervor, dass dies ein sehr seltner Wurm 
sein muss. Hier muss ich noch bemerken, dass, dem Kopfe nach, 
dieser Wurm der Taenia filicollis Rud. nicht unahnlich ist, sich 
aber von ihr durch die langeren Glieder, wie wir es unten sehen 
werden, unt erscheidet . 

Die Lange des grossten Exemplares betrug 45 Mm. bei 
einer Breite von 1 Mm. Der Kopf ist verdickt und endigt mit einer 
Spitze der keine Hacken tragt; die 4 Saugnapfe liegen paarweise 
auf der Grundhalfte des Kopfes; sie sind etwas in die Q.uere ver- 
langert und stark muskulos, v/eshalb der Wurm auch so fest an die 
Wande des Darms sich ansaugt, dass men beim Einsammeln sehr vor- 
sichtig sein muss, wenn man nicht "Fragmenta acephala" erhalten 
will. Das zu seiner Basis verengte Kopfchen geht in den ziemlich 
dunnen Hals uber, welcher ungefahr 12 Mm. lang und 0.5 Mm. breit 
ist. Hinter dem Halse fangt die Kette an, in der ich 23 einzelne 


Glieder gezahlt habe. Lie ersten, resp. jungsten olieder sind 
fast vollkommen quadrat isch; weiterhin werden sie etwas langer, so 
dass die letzten acht Glieder 1,4 bis 1,5 Mm. lang sind; dabei 
betragt ihre Ereite 1 Mm., beiji einer Dicke von nur 0,3 Mm., so 
dass der Wurm vollkommen plattgedruckt erscheint . Das letzte Glied 
ist immer von hinten abgerundet. Dies ist alles was wir an einem 
unladirten Exemplare zu sehen bekamen. 

Wenn wir nun aber einzelne zuvor mit Cariiin und Glycerin 
bearbeitete Glieder und dunne Querschnitt e unter dem Mikroskop un- 
tersuchen, so uberzeugen wir uns, dass der Bau der Korpermasse 
dieses Bandwunnes sich uberhaupt durch nichts von dem Bau anderer 
Bandwurmer unt erscheidet . Wir finden hier wie auch bei Cyatho- trUncatus, dass der Kcrper aus folgenden Elementen besteht:- 
aus der kornchenreichen Grundsubst anz , die von sich schlangelnden 
dors-vent ral en Muskelbundeln durcheetzt wird; einer Schicht L'angen- 
muskeln und einer Schicht Quer- Oder Ringmuskeln, einer Schicht 
feinkorniger Rindensubstanz , in der die Enden der dorso-ventral ■ n 
Muskeln liegen, und endlich der sehr f einer ausseren Haut , die gar 
keine Muskeln besitzt. In der Langszxe eines jeden reifen Gliedes 
liegt das centrale Rohr des Eierstocks, welcher aber manchmal naher 
zur einer Oder andern flachen Seite zu liegen kommt , wie man es 
auf den Querschnitten bemerkt. Von diesem Centralrohr verlaufen 
nach beiden Seiten zu unregelmassig gebogene und theils sich auch 
verzweigende Auslaufer, die in ihrer Large Mittelst der dorse- 
vent ralen Muskeln erhalten werden. In der Mitte des Hauptrohres 


geht von ihm die Scheide ab, die, nachdem sie eine schwache .Biegung 
gemacht hat, sich in der Mitte der seitlichen Oberflache des Gliedesj 
nach. aussen (iff net; die Scheide ist ziemlich breit , so dass die 
Eier sie leicht passiren konnen. In dem von mir untersuchten Exemp- 
lare waren die Eier in der Mehrzahl der Glieder vollkommen reif;- 
sie enthielten eine grob und- feinko'rnige Masse und waren mit 
einer ziemlich dicken Haut utgeben. Aben wie in diesen reifen Glie- 
dern, so auch in den jungern bin ich nicht im Stande gewesen die 
sog. Dotterstocke aufzufinden; ob sie hier so fruh verschwinden , 
Oder zu fein sind, weiss ich nicht; auf den Q.uerschnitten sind 
sie aber nicht aufzufinden. 

Gleich uber der Scheide liegt und offnet sich nach 
aussen das mannliche Zeugungsorgan ; dasselbe besteht aus einem 
ziemlich grossen und kolbenf ormigen Cirrusbeut el , in dem der einge- 
zogene Cirnis liegt; der Cirrus ist ziemlich dick und am Grunde 
dunner als am Bnde; dessen Kanal ist durch. den Cirrusbeutel zu 
sehen. In den Warden des Cirrusbeutels trafen wir, wie immer, 
Langs- und Quermuskeln. Die Sa^.endrusen waren nicht zu sehen. 

Ties ist alles was ich von dem Bau des Bandwurmes erf- 
ahren konnte. Die Unvallkommenheit meiner Untersuchung wurde aber 
durch den Mangel an Material bedingt. Dennoch aber fuhle ich mich 
berechtigt zu sagen, dass auch dies Wenige genugt , urn dem besch- 
riebenen Bandwu.rm seine haturliche Stellung in der Reihe seines 
Gleichen anzuweisen und somit die Zahl der zweif elhaf t en Art en 
su beschranken. 

TToch muss ich hinzusetzen, dass aus dem Gesagten doch 


dlar ist, class ich wchl mich nicht irre, indem ich den von mir 
auf gefundenen Wurm als dem Bothriocephalus barbatulae Rud . iden- 
tisch halte. 

While the foregoing description is sufficient to warrant 

the inclusion of the species in the genus Prot eocephalu s it is not 


sufficient to enable one to determine relations of the species with 
the other members of the genus. In size the fcrm resembles P. 
gaster ost ei and P. agcni s . "Per i-he present the species must be put 
in the list of incompletely described species. 


(PI.? Figs.)72-/7^ 
1906: Icthyotaenia sulcata Klaptocz, 1906:123-130. 

SPECIFIC DIAGNOSIS.- The characters of the genus. Cestode 


of small size, up to 68 mm long. Max. breadth 1.5-2.0 mm. Scolex, 
very variable in form, unarmed, without apical depression or fifth 
sucker, divided by distinct furrows into four quadrants. Median 
furrows arise near first proglottids. Cross-section of one type of 
scolex nearly round, of the other elliptical. Breadth of scolex 0.46 
1.75 mm, thickness of same 0.67-1.28 mm. depending on contraction 
states. Suckers, deep rounded, one in each quadrant. Diameter of 
sucker 0.250 mm. 

Neck none. First proglottids "broader than long. Mature 
and ripe proglottids quadrate or longer than "broad. Last proglottid 
rounded posteriorly. Max. breadth of proglottid 1.5 mm. Max . 
length of same 2.0 mm. 

GENITAL ORGANS.- Genital pore at end of first third of 
segment. Cirrus-pouch about 0.200-0.250 mm long, extending about 
; l/4 across the proglottid. Several coils of ductus ejaculatorius 
in cirrus-poach. Cirrus when protruded long and slender. Testes, 
about 200, irregularly arranged between vitellaria. 

Opening of Vagina posterior to the cirrus-pouch. Lumen of 
vagina large. Ovary, posterior, biloted. Vitellaria, follicular, 
lateral. Uterus with 10-12 lateral out-pocket ings in ripe proglot- 
tids. Eggs, pear-shaped, circular in cross-section. Diameter of 
same 0.016 mm, length up to 0.019 mm. 


HABITAT.- Polypterus Endlicheri Heckel, taken at Due/a on 
the White Nile, and Clarotes la t iceps R uppell, from the White Nile 
in the region of Khartcum. 

The material was collected by Dr. F. Werner in the Spring 

of 1905 and was described "by Klaptccz (1906). In the summary and 

also in the following description the data are taken frc?- 

Klaptocz* paper and from his drawings. 

EXTERNAL CHARACTERS . - A specimen from Clarotes measured 

63 ram .one from Pclyterus 60 mm. 

The scolex is of very variable form, is unarmed, has no 
apical depression, and it always shows four very distinct furrows 
of which the two on the median line are the heavier. These divide 
the anterior face of the scolex into four similar large quadrants 
whose acute angles come together at the scolex tip. The free 
margin of the quajftdrants eft en are indented especially on the 
flattened sides. The furrows, expecially the heavier median ones, 
arise in the region of the first proglottids. 

In each quadrant a deep rounded sucker is sunken, which 
lies nearer the scolex margin than to the scolex tip. 

Twc types of scclex are recognized with intermediate 

[fluffy ni) 

forms. The one^is more or less flattened on the anterior face. It 

has a mere or less round cress section. Its furrows are shallow, 

and its suckers are directed anteriad //bile the other type. has 


an elliptical transverse section. Its thickness is less than the 
breadth of the scolex. The scolex itself is much flattened. The 
furrows are deep and very plain, reaching nearly to the tip of the 


scolex. The suckers are more slongated and the sucker openings are 

directed outward. These are to "be considered contraction states of 

the same species since int enr.ediate stages are present. The second 

type is found in Po lypterus , and the first type in Clarotes . 

The diameter of the scolex in specimens from Clarotes 

lat iceps is up tc 1.72 mm, the diameter of a sucker of the same 

0.250 mm. The diameter of the smallest scolex of this host is 0.75 


mm. The thickness of scol ices^ Polypterus Endlicheri ranges from 
0.67-1.28 mm, the "breadth from 0.46-675 ram^ ( a remarkable variation 

Ko neck is present. The youngest proglottids are thinner 
than the head and only slightly narrower. Their breadth exceeds 
the length. Proglottids increase in both breadth and length with 
age until they are quadrate in form or longer than broad. The last 
proglottid is rounded posteriorly. The max. "breadth of a proglottid 
from Claret e3 is 1.5 mm, max. length 2 mm. The max. breadth of a 
segment from Polypte rus is 1.5 mm. and the max. length of the same 
1 . 9 mm . 

Sexual ripeness appears very early. The anlage/i of genital 
organs in stained specimens may be seen in very young proglottids 
within two scolex lengths of the head. Many proglottids are sexu- 
ally ripe before becoming quadrate in form. 

GENITAL DHGrAKTS.- The marginal genital pore alternates 

irregularly. Sometimes in as many as ten to twelve or more 

proglottids the genital sinus may be on the same side^tho this con- 
is trup 

dition,^ rarely t c in more than five segments. The sinus is situated 


at about the end of the first third cf A segment. The vagina opens 
posterior cc the cirrus- pouch. 


Male organs .- The cirrus-sheathe^containing the coiled 
cirrus is of a pear-shaped form and lies perpendicular to the margin 
of the proglottid. It may extend l/4 across the "breadth of the 
proglottid. The cirrus-pouch is about 0.200-0.250 mm. long (meas- 
ured from Kiaptocz' figures which are drawn to scale). 

The cirrus when protruded is very long and slender. It 
is thicker at the base than at the point and is devoid of hooks or 

The testes are elliptical, in number, about 200. The 
testicular field is limited by the vitellaria, the anterierr margin 
of the segment and the ovaries. There is no median zone free from 
them except where they are pushed aside by the . developing uterus. 
As the uterus continues to develop tne testes are pushed well into 
the lateral fields. i 

Fe male org ans.- The vagina. which has a large lumen runs 
to the middle of the proglottid in a mere or less sinuous course, 
then bends ana passes directly oack to the posterior end of the 
segment. In proglottids with developed uterus the vagina, is 
pushed somewhat from the median line, and, it seems, always to- 
ward the side on which it opens. 

The ovary is posterior, is hilobed, and the lobes are 
limited, laterally by tne vitellaria. In the end proglottid the 
lobes of the ovary are bent together. 

The vitellaria are follicular and lateral, reaching from 
end to end of the segment. On the atrial .side the vitellaria is 
divided into two parts which are connected by small follicles which 



lie docrsal ard ventral to the genital passages. 

(PA f f*T l7 f 

The uterus^ is a median tube reaching to the anterior end 
of the proglottid from which arise many (10-12 in drawing) lateral 
out pocket ings . These are separated when fully developed by thin 
t issue-layers. 

The eggs are pear-shaped, circular in cross-section, 0.016 
mm. in diameter and 0.029 mm. long. 



(PI. 7 Fig,/^) 

1877: Taeni a c;, clops ^ on Lin stow, 1877:15 
1905': Proteocephaius iongicolli s Nufer 190£ 152, in part, 
von Linstow's description of this immature form is here 


TAENIA CYCLOPS n. sp.« In Coregonus maraena aus dem Schall- 
See. Die Tanie ist 25 Mm. lang und 0,27 Mm. breit , uberall fast 
gleich "breit, die letzte Proglottide hinten zugespitzt . Die Kalk- 
kcrperchen sind klein, ohne concentrische Schichtung; das Kopfende 
1st abge rundet , die Saugnapfe sind langscval, 0,15 Mm. lang und 
nach hinten zugespitzt, 0,1 Mm. "breit; ein funfter scheit elstandiger 
Saugnapf hat 0,069 Mm. im Durchmesser ; Geschlecht sorgane waren noch 
nicht vcrhanden. Taenia lcngiccllis aus verschi edenen zur Familie 
der Lachss gehorenden Fischen hat zum Unterschied vcn disser Art 
langlichrunde Saugnapfe, deren langerer Durchmesser rechtvdnklig 
zur Langsaxe des Thieres steht. 

ITufer (1905) thought that this form was the larval state 
of P. longicOll is (Pud.) however the character of the suckers do not 
permit of such a determination. P. cylops is remarkable for the 
notch in the inferior margin of the sucker. This form must be 
considered as a species inquirenda. 


1859: Taenia h emisphaerica Molin 1859:14. 
1896: I chthyo taenia hemisphaeria Riggenbach 1896. 

Molin' s original description and observations are as 


TAENIA HEM I SPHAERICA . - Caput heini Bphaericum , acetabulis 
ant icis ; ore inermi ; collum longissimum . ant ice dilatatum ; articuli 
supremi brevi ssimi , subsequent es transverse pa ra il elogrammici , 
post remi subquadrat i ; aj. erturae genita,les marginales, vage alt erna e . 
Longit . 0.07 ; lat . 0.005. 

HABITACULUM, Anguilla vulgari s : in intestino tenui, 
Movembri, Patavii (Molin.) 

OSSERVAZIOUE 1. Ai 5 di Novembre del 1858 rirvenni nell' 

intestino di un* anguilla, nella quale avevo trcvato 5 Dibothrium 

clav i ceps , un esemplare di quel verme; ed in un aJra anguilla un' 


esemplare intero e varii frammenti. 

OSSSRVAZIOUE 2. Fegli articcli bene sviluppati potei 
distinguere ai margini laterali le ovaja, e tutto il resto dell' 
articclo riempito call' amplissimo ovidotto ravvcltc in ambagi e 
ripieno di uova sferiche. 11 el mezzo vidi collocate orizontalmente 
il m.embrc virile attoriglioto che si estendeva fino al centre dell* 
j articolo. 

OSSERVAZIOKE 3. Fino ad ore: non si conosceva dell anguilla 
altro che la Taenia macrocephal a , eel la quale pert la hemisphaeri ca 
si distingue per la forma della testa ncn che per la lunghezza del 


collo. Questa N e affir.e alia f ili colli s , dalla quale perb si dis- 
tingue per la forma della testa e del colic, e deve percio venir 
regi strata dcpc di cuesta nel sistema. 

Since the appearance of Molin's description no further 
attempts have been made to describe this species. In fact I have 
been unable to find a further notice of its having been found. 

Linton (1886) thought that his (Taenia) P rot eoc ephalus dilatatus 


might be the same as ( Taenia ) P. hemisphae ricus . Riggenbach (1896) 

— — — — — f 

said, M Ein genauer Vetgleich der Diagnosen, welche die genannten 
Aa:oren den fraglichen Species gaben, zwingt mich mit Bestimmtheit 
eine I dent it at der I ( chth^p taeni a) dilatata Linton mit iZX cht hyo- 
taenia ) h em ispha erica Molin anzunehmen." 

Schneider (1903) identified a specimen from Anguilla 
vulgaris as . Ichthyo taenia hemisphaerica . Later (1905) he de- 
cided that this form was not 3C, hemi sphaerica but I. macrocephala 
(see the discussion in the historical summary of P. map rop- e phalus ) . 

Nuf er (1905) attempted to show that P. dilatatus Linton 
and P. hemispha e ricus Molin were identical, and further that these 
species were identical with P. mac r oc ephalus Creplin. 

In the discussion of P. mac roc ephalus in another part of 
this article I have shown that P. dilatatu s Linton and P. 
mac r co ephalus Creplin are identical. I am not sure that P. 
macrcpephalus and P. hemisphaericus are identical. 


1867: Taenia micrcpteri Leidy, 1887:23. 

Leidy 1 s (1887:23) description is as follows: 
^TAENIA 1OCP0PTEFI . - Head large, compressed spheroidal, 
with four subterminal spherical bothria and a papillifonn unarmed 
summit; neck none; body obscurely segmented, and with no obvicus 
internal orga-is, posteriorly variably narrowed and obtusely rounded 
at the end. Length from half an inch to an inch, and about 1 mm. 
wide. Apparently a larval form; fcund in the body cavity of the 
Black Bass, Lti crept erus nigricans . Six worms, soft, white, and 
active. The longer ones of an inch would elongate to double the 
length becoming proportionately narrower. The head, about 1 mm. or 
more in miameter, varied in length and breadth, according to 
contraction, sometimes one and sometimes the other being the larger. 
Lake George, N. Y. 5 

It seems highly probable that this is a larval form cf 
P. ambloplit i s since Leidy originally described that species from 
the Ambloplit eg rupestris of the same lake. Moreover P. amblopl iti g 
has been found in Micropt eru s dolcmi-n and M. salmoides as well as 
in several bther host species. No other cestode species than P. 
ambloplit is and this larval P. micropt eri have been reported from th 
Cent rarchidae of U. America.Leidy 1 s record is not accompanied by 
drawings, his specimens are not available, indeed, they may never 
have been preserved, hence a positive determination is now impossibl 



(Pl.<? Fig4#) 
1887: Taenia simplici ssima Leidy, 1887:22. 

Leidy (1887:22) describes this form thus: 
U TAEFIA SIMPLI CI SSIMA. (drawing).- Hea.d small, unarmed, 
true ate; bothria spherical, terminal, occupying the four angles; 
neck very long, nearly or as wide as the head, "body gradually widen- 
ing to the posterior third and then tapering; anterior segments 
transversely linear, subsequently reversed disklike, gradually 
longer and wider, then campanulate and gradually becoming longer 
and narrower. Generative apertures and ova unobserved. A number of 
specimens from the Cod, Gadus cal larias , up to 20 lines by 1 mm. 
where widest. Two only of the specimens retained the head. (Drawing] 

This species was induced in a list of species of 

P,~Y o t e a c e phal u s feiven by Riggenbach (1896:267). However he made 

no comments on its structure or position. It seems quite improbable 

that this form is a species of FCrot e o c e pha l u s . My chief reason 

for so thinking is that the host, Gqcus c allarias , is a purely 

marine fish, and thus far no unquestioned species of P Crot e s cephalus 

has been taken from a strictly marine fish. The infection of 


Anguil la chry sypa A. vulgaris with F, ma c roc ephalus . probably occurs 
while the eel is in its fresh- water habitat. In Riggenbach 1 s 
(1896:246-248) list of hosts and their parasites are two host 
species, Bel one acus and Merlangus pcllachitis which live in salt 
water. Beldne acus is credited with the parasite, Ichth yotaenia 
belone s Muller, which Mcnticeili (l82»i) thought was very probably 


identical with Bothriocephalu s belones Dujardin. Merlan gus 
gollachius is credited with ICchthy ot aenia. pollachii which Monticelli 
(1891) thought was probably identical with Abothrium Gadi van 
Beneden . 

With the data tc be gleaned from Leidy's drawings^ which 
are reproduced (PI fj f igjri , a positive determination is impossible. 


(Pi. 7 Ji&.}7?,t?fJ 
1897 Taenia salvslini Linton, 1897:424. 

Linton (1897) described this species thus: 

<( TAENIA SALVELINI . Type .- Fo. 4811, U.S.N.M. From intestine 
of Great Lake trout ( Crist ivomer namaycush ); Outer Island, Lake 
Superior; J. W. Milner. In this lot are several small Taeniae ; 
scolices with few roundish or oblong segments, in some of which the 
male genitalia were developed, otherwise immature; length from 3 
to 5 mm. 

These specimens "bear some resemblance to the young of T_. 

t orulos a, Batch, as described by Zschokke. (1882-28). 

Head usually rounded in front, with suckers directed 
anteriorly. The diameter of the head varies according to state of 
contraction, usually 0.3 to 0.4 mm.; suckers relatively large, 
usually directed forward, prominent, rather deep, 0.15 to 0.18 mm. 
in diameter, aperture 0.1 mm. in diameter. ITeck assuming various 
contraction shapes; in some continuous with head and merging im- 
perceptibly in body; in others separated by a constriction behind 
the head; in others not separated from head by constriction but 
with segments beginning abruptly; numerous calcareous bodies in 
neck behind head. Usually first segments indistinct, broader than 
long. Oldest segments longer than broad; genital aperture near 
middle of lateral margin a little nearer anterior margin. Yitellaria 


making a narrow border along lateral margins; beginning of germ 
gland at posterior end; central region in front of rudimentary 
germ gland occupied by prominent testicular lobes; cirrus-bulb 
Oolong, elliptical, convex on posterior, straight or slightly con- 
cave on anterior surface; cirrus enters from inner anterior edge of 
bulb, then traverses middle of bulb to orifice; length of bulb, 
0.26 mm.; diameter, 0.14 mm; largest segments, 1.5 mm. in length 
and 0.7 5 mm. in breadth.'* 

Since this description is insufficient for determination 
in comparison with the forms described here^Dr. F. B. Ward very 
kindly secured for me Linton's specimens from the National Museum 
at Washington. Unfortunately but fragments remained. These 
fragments were cleared and examined in glycerine. My observations 
on the heads agree very well with Linton's records. I am unable to 
find a fifth sucker or a trace of an end organ. The suckers in 
their maximum diameter measure 0.170 mm. the sucker opening about 
0.100 mm. The head is flattened dorsovent rally. In general shape 
it resembles some dilated heads of P. ping r- ls n . sp, but is larger. 
Linton says that it resembles young specimens of P.torulosus, how- 
ever its measurements are quite different. It must be left for the 
present in the list of inadequatly described species. 


(PI. q Yir.3.)*f,l* c ) 
1904: Ichthyotaenia skorikowi , v. Linstow. 1904; 18-19 


From Von LinstcW's (1904) description the following is 
taken: Length up to 200 mm; breadth anteriorly 1.14 mm. First 
proglottids beginning directly behind scolex very short. Breadth 
in middle 3.75 mm; length of proglottids here 0.95 mm. Breadth at 
posterior end 3.16 mm; length of proglottids here 1.97 mm. Last 
proglottid rounded posteriorly. 

Scolex short, 0.67 mm. broad. Suckers 0.250 mm, in 
diameter with a fifth apical sucker 0.130 mm. in diameter. Muscles 
of subcu .ticular layer as usual. The heavy layer of longitudinal 
muscles below the subeu ticula anteriorly are very strongly devel- 
oped, filling up almost the entire space in unripe proglottids. 
Dor so-ventral muscles very heavily developed The- main longitudinal 
i nerves in anterior segments lie in the 12- and 88 hundredths of the 
segment-width in ripe segments in the 6-and 94 hundredths* Within 
the nerve trunks lie the excretory ducts two on either side. 
Chalkbodies are lacking entirely. 

Sex openings are marginal, irregularly alternating, 
posterior to the middle in each proglottid. The vagina lies an- 
terior to the cirrus. The cirrus is con©- shaped, blunt at the tip, 
0.141 mm. long. The point is 0.042 mm. broad. The cirrus- sheath 
extends l/5— l/6 across the breadth of the proglottid. The vas 
deferens makes many coils before entering the cirrus-sheath . 


Testes are very numerous, 0.053-0.083 mm. in diameter. 

Vagina runs in a curve to the middle of the posterior 
region of the segment where it ferns many coils a.nterior to the 
ovaries. These coils function as a receptaculum seminis. The 
ovary is made up of two groups of club-shaped bags which run parallel 
to the posterior margin of the proglottid. Anterior to its middle 
lies an oceapt. Vitellaria are f ollioular^reaching on left and 
right the full length of the proglottid inside the longitudinal 
muscles. The uterus in the posterior part fills the whole space 
within the longitudinal muscle fibers and extends arteriad in round 
diverticula. In his drawing there are 6-8 diverticula on either 
side. His drawing is reproduced (PI. f fig. I8£>). 

EGGS: Globular, 0.027 mm. They have two shells, the 
outer very delicate and hyaline. The six-hccked embryo measures 
0.021-0.023 mm. in diameter 

HABITAT: Intestine of Acipenser st e llatus , River Gurgen, 
Caspian Sea. 

v. Linstcw failed to state who collected the specimens of 
this species and the abundance of their occurrence. 


1888: Taeni a leptosoma Leidy, 1888: (nec Diesing), 

1890: Taenia nematosoma Leidy, 1890. 
1891: Taenia somatol epta Mont i c ell i, 1891. 

In May 1888 Leidy reported on the "Parasites of the 
Pickerel" as follows: - 

PARASITES OF THE PICKEEEL.- Dr. Leidy remarked that among 
the numerous parasites which are rientioned as infesting the Eike, 
Esox luci us, of Europe, no Taenia is indicated. In the Pickerel, 
Esox ret iculatus , brought to cur market, a species of the latter 
appears to he common. In two fishes he found half a dozen in the 
intestine and stomach; and in another a single individual two feet 
in length. It resembles closely the Taenia anblo^ lit is, noticed in 
the Rock Bass, Ambloplitis rupes t ris (Proc., 1887, 23), and may be 
the same. Distinguishing it with, the name of Taenia leptosoma, its 
characters are as follows: Body long and thin, and at the forepart 
thread-like. Head unarmed, without rostellum, with four equidistant 
hemispherical bothria; neck very short or none; anterior segments 
transversely linear, many times wider than long; posterior segments 
gradually becoming proportionately longer and quadrate and barrel 
shaped; genital apertures marginal , alternating irregularly. Ova 
spherical . 

Length from 6 to 9 and 26 inches, shortening tc one half 
or less; breadth to 2 and 2.5 mm. Head 0.25 to 0.5 mm. broad; 
bothria 0.125 to 0.175 mm. Anterior segments an inch from the 
head 0.175 mm. long by 1 mm. broad; posterior segments 0.5 to 0.75 


mm. long by 2 to 2.5 mm. "broad. Ova 0.028 to 0.032 mm. in diameter. 

A single slender Scolex associated with the longest Taenia 
was 4 mm. long "by 0.25 wide, but elongated to 8 mm . by 0.1 wide. 
The head was of the same form as that of the Taenia. After being 
in alcohol, the head of the Scolex was 0.225 mm. wide, with the 
bothria 0.1 in diameter. The posterior part of the "body exhibited 
traces of segmentation, with the segments 0.075 mm. long by 0.25 

In November 1890 from the same host species Leidy re- 
ported Taenia nematosoma. J quote his description! 


Taen ia nemat osoma, n.s.- Head rounded quadrate, unarmed, 
with equidistant hemispherical bothria and a small central papilla; 
neck short or none; fore part of body linear; anterior segments 
much wider than long; posterior segments gradually becoming pro- 
i portionately longer, quadrate or barrel- shaped; genital apertures 
marginal and alternating irregularly. Length to 9 inches, contract- 
ing to about one-half. Breadth of head 0575 to 0.5 mm.; bothria 
0.175 wide; neck 0.25 v/ide; anterior segment an inch from the head 
0.175 mm. long by 2 mm. broad; posterior segments 0.75 mm, long by 
I 2 mm. broad, and when contracted widening to 2.5 mm. Ova spherical, 
| 0.028 to 0.032 mm. 

A half dozen specimens in the stomach of two Pickerel, 
Esox reticulatus . 

Neither report was accompanied by drawings. 


Monticelli (1891) on account of the confusion of this 
name with Taenia leptc soma Diesing suggested the name Taen ia 
somatolept a . However the name nematosomus sh ould be prefered 
because it is an earlier available name than gomat clep tus . 

Leidys tv/o forms agree in practically every particular 
as to measurements and proportions and host species. I therefore 
conclude that they are identical and I include them both under the 
name P.^ematoso mus (Leidy) . This species differs from P. ambloplitis 
chief ly in the measurement of the suckers. Its measurements vary 
greatly from those of P. pingujs n.sp. and from P. esocis Gui 
Schneider. None of Leidy's specimens remain so the species must 
continue a species inquirenda. 


CHOANOSCOLSX ABSCISUS. ( Riggenbach ; 1896 ) La Rue. 

(pi. °{ Fig. (S3 ) 


189 G: Ichth; taenia absoisa Riggenbach, 18961 193-210 
SPECIFIC DIAGIT03I3.- Characters of genus. Worms small, 
25-30 mm. long, breadth up to .765 mm. Number of proglottids up t 
70. Head somewhat conical^ a fold of tissue at the base at times 
partly covering the suckers. Apex of head flattened. Max. breadth 
of head 0.476 mm.- No rostellum, no fifth sucker. Suckers large, 
oval , 0.3o6 mm. long x 0.136 mm. broad. First proglottid longer 
than broad. Tenth proglottid 0.476 mm. broad x 0.102 mm. long. 
Segments near middle of worm broader than long or nearly quadrate. 
Last proglottid : .02 mm. long x 0.765 mm. broad. Proglottid 
margins straight j posterior corners rounded. Last proglottid with 
I bluntly rounded end. 

Sex organs.- As in genus. Genital aperture situated 
at end of anterior 1/3-1/5 of the proglottid. ITo genital papilla. 
Testes about 100 in number, 0.054 x 0.045 mm. in size, arranged in 
a single layer between vitellaria. Coils of vas deferens voluminou 
Coils of ductus e jaculatorius few. Cirrus swollen, with large 
lumen. Cirrus-pou h large, pear-shaped, muscular, reaching nearly t 
middle of segment. Length of cirrus-pouch 0.34 mm. 


Vagina, anterior or posterior to cirrus-pouch. Lumen of first part 
of vagina large. Keceptaculum seminis present. Ovary bilobed, 
posterior, unbranched Vitellaria, lateral, follicular. Uterus a 
median tube with many lateral outpocketings. Eggs. 0.021 mm. long 
x 0.016 mm. "broad. 

5ABITAT.- Silurus sp. Rio Paraguay, Gouth America. 

The material was collected "by Dr . jernets in Paraguay, 
1894. The description is based on the work of Hig^enhach (1895). 
Riggenbach considered this form to be a species of I-chthyotaenia 
and his comparisons are made with species of that genus rathor 
than with Coral: obotlirium. 

EZTERITA1 CHAl-.ACTERS.- The worm is small being 25-30 mm. 
in length. It is made up of about 70 proglottids whose form.varie 
ac or ding to the age. 

The scolex varies from the usual shape in being some- 
what conical instead of spherical and in having at the base of the 
cone a fold of tissue or a mantle which may be partially drawn 
over the suckers. The tip of the cone is flattened. Jhe 

maximum breadth of the head is 0.476 mm. and the breadth of the 


flattened apex of the cone is : .255 mm. 

The suckers which are very la^ge in pre port ion to the 
size of the head, are separated from each other by only a small 
zone. The shape is oval, at times triangular. Their length is 
about 0.306 mm. and their breadth about 0.136 mm. 


The head reaches its greatest breadth, 0.476 mm, at the point 
where' it passes over into the neck, which has a breadth of 0.348 
mm. Tho neck is of mediarn length. 

The first proglottids are very narrow transverse bands. 
The tenth proglottid is about 0.476 mm. broad by 0.102 mm. long. 
The length of the proglottid increases with age more rapidly than 
the breadth. The twentieth proglottid has a length of 0.153 mm. 
j Proglottids near the middle of the worm are broader than long or 
quadrate. The last proglottid is 0.765 mm. broad by 1.02 mm. ] ong. 

The proglottid margins are straight and the posterior 
corners of the proglottids ar rounded. Except for the slight 
raising of the margin in the region of the genital opening, the 
geometrical lorm of the proglottid is not disturbed. The last 
proglottid alone makes an exception, for in this case the end 
of the proglottid is rounded off. 

IiTT;;H 'AL Ai-Ai'fHY.- For the description of the euticula, 
i arenchyma, musculature, and nervous system see Riggenbach (1895J 
196-199). These structure are typical for Proteocephalids. 
The musculature of the head Rig-enbach did not describe. 

Excretory system .- In the head the small space between 
the suckers is almost completely filled with vessels of the 
excretory system. 


The form main vessels are bent inward as they enter the head. A 

circular anastomosis "behind the muckers could not he demonstrated. 

In the short apical region of the head the Teasels are never 

capillaries and a capillary apical plexus such as is found in 

Corallobotj ium 1 obo 3 am does not occur. Eiggenbach was not able 

to find that the vessels of the head or of the plexus were in 

direct connection with the exterior. 

Posterior to the suckers fhe main excretory vessels pass 

to the lateral fields of the neck in a course that is perpendicular 

to the longitudinal axis of the bodjr, then they bend posteriad, 

decrease in size and extend throughout the strobila no arly A straight 

narrow tubes, the dorsal lying above the ventral tube. In the 


region of the cirrus-pouch vagina the dorsal vessel passes above and 

^ventral vessel below these organs. The excretory vessels lie 
mesad of the vitellaria and the lateral nerve, trunks. At the posterior 
part of each segment the main excretory vessels are connected by a 
transverse anastomosis . 

In the region of the transverse anastomosis are canals 
which connect the main vessels v/ith the exterior . These canals 
arise directly from the main vessels and do not arise from a 
capillary network. The point of opening to the exterior is on the 
surface near the posterior angle of the proglottid. 

In the posterior part of the end-oroglottid the four main 
vessels join together to form a small blade er or reservoir. This 
reservoir opei. 3 in the mid d le of the rounded posterior margin of the 
end-progiott id. 


remale organs .- The vagina opens into the common genital 
sinus anterior or posterior to the cirrus-pouch. In old oroglottids 
the beginning part of the vagina is swollen into a sac nearly as 
large as the cirrus-pouch hut in .young proglottid?, the diameter of the 
vagina is uniform^ In its course to the middle ol the proglottid it 
describes an ace, thon it bends sharply and passes to the inter- 
ovarial space in a spiral or sinuous course. The vagina crosses the 
convolutions of vas deferens only when it opens anterior to the cirrus 

A receptaculum seminis is present near the middle of the 

ovary. This is more than a broadening of the vagina at this point 

for in liiggenbach 1 s drawing there is shown at this place a change 

in the histological structure of the vagina. The relations of the 

organs v/ithin the interovarial tfpace is similar 

of Proteocephalus 
to that in species ana need not be discussed here. 

The bilobed ova^y is in the pos J erior region of the 


proglottid. The lobes are unbranched. sacliko structure, united by 

a mi&piece from which the oviduct arises. 

The vitelline glands are lateral follicular structures 
extending the ful^Length of the segment. They occupy a broad zone 
outs:'de of the excretory vessels and the nerve trunks. 

The uterus is a median longitudinal tube with a large 
number of lateral outpocketings on either side. The uterine eggs are 
0.021 mm. long x 0.016 mm. broad. The elongated shell is very thin 
and surrounds in part the yet undifferentiated egg-coll with the yolk 



cells j in part the cell-groups which are the results of the first 
developmental stages? Evidently Riggenbach saw no embryos. 

SYSTEMATIC I OSITION.- In life the heads of typical species 
of Proteocophalus are extremely variable in form passing with con- 
siderable rapidity from one contraction state to another. By the 
contraction of longitudinal muscles extending into the apex of the 
head the suckers may be drawn down out of sight within an 
inflated neck region^but there are no folds of tissue at the base 
of the head within which the head may retreat. 

In the species just described this fold of tissue with^in 
which the suckers are partially withdrawn seems to be a constant 
feature of the scolex. Thus the head differs from those of species 
of Proteocephalus *■ 

At first one notes certain similarities with the 

of species of CoraqiobotT irium further consideration Oi the structure 

of the two types of heads convinces one that they are not alike. The 

the suckers are directed anteriad 
head of a Gorallobothium is greatly flattened anteriorly and^maygiac 
At the margins 

of the flattened apical region are numerous folds and lappets which 

form a corolla-like sheathe- about the suckers* In this soecies the 

In CoroiLlobothrium lobosum 
head is conical and the suckers are directed outward. J Kiggenbaoh 

(1896) found a small muscle-cross connecting each dorsal sucker with 
the ventral sucker opoosite it. Riggenoaeh did not see such a str 1 
iture in his sctions of the headsof I ohthyotaenia abocisa . It is 
doubtful ii it occurs in any other genus known at present to be- 
long in this family. 


One may consider that I cththyo taenia abscisa Riggenbach 

forms a transitional form between the genus Proteocephalus on the 

one band and G o r oj 1 o b o tb riumi on the other. This belief finds its 

support in the fact that the heads of the pier oeercoids of an 

unknown species of Corallobothriurn found encysted in the liver of 

Ame iu rus m e lac and nebulosus from the Illinois river much 

resembles the head of I c h t hy o t a e n i a abscisa Riggenbach. In these 

plerocercoids, the folds and lappets about the head v/ere much less 

completely developed than in the adults* 

The heads of living plerocercoids were somewhat conical, 

not flattened on the apex as in preserved heads of the adult worms. 

There is a marked difference, however, between the heads of these 

plerocercoids and the heads of Ichthyota e nia abscisus • In these 

plerocercf ids the suckers are plainly p.'ired, two dorsal and two 

ventral, while in I. abscisa the head is but slightly flattened 

dorsoventrally and the suckers are not plainly paired. 

Since I chthyotaenia abs cisa d ee not come within the genera 

Proteocepha lus or Goro llobothrium I propose for it the new °:enus 
The type species in 

UnoanogcoleX j t^scisu ^ (Riggenbach 1396) La Hue 1911, from Silr.rus 

sp, Rio Paraguay, South America. 



fpl. t :±r-. H% IX< °) 

SPECIFIC DIAGNOSIS.- Characters oi the genus. Length 
up to 36.0 mm. Max "breadth about 2.0 mm. Head conical, flattened, 
divided by grooves into four quadrants. Breadth of head 0.g55- 
0.403 mm., thickness of same up to 0.506 mm. length up to 6.270 mm. 
Suckers four, circular, oval, or at times somewhat triangular , 0.105 
0.170 mm. in max dimension. Cavity of sucker 0.055-0.105 mm in 
diameter . 

ITeck long, 5-7 mm. in length "by 0. 170-0. 4£5 mm. broad. 
First proglottids short. Mature proglottids quadrate, 2.0 mm. in 
length and breadth, or somewhat longer than broad. Ripe proglottids 
measure up to 3.0 mm. long by 1.2 mm. broad. 

Segmentation indistinct. Proglottids attached by full 
Width. Surface smooth. In life and when preserved specimens some- 
what translucent. 

GENITAL ORGANS.- As in genus. Genital pore marginal, 
irregularly alternating, situated near middle or at end of first 
one third of proglottid. Vagina anterior or posterior to cirrus- 

Male Organs .- Teste 1 0-215, oval o. polygonal measur- 
ing up to 0.053-0.106 mm. in two fields Vas deferens in ripe 
proglottids a heavy mess of coils reaching from end of Cirrus-pouch 
to the mid-field. 


Cirrus-pouch 0.255-0.320 mm. long by 0.080-0.090 mm. "broad. Ratio 
length of cirrus-pouch to proglottid "breadth from 1:4 to 153. 
Cirrus slender, about 0.20 mm. long when protruded. 

Femal e organs . - Vagina dilated in first part, not cross- 
ing cirrus-pouch. Sphincter vaginae j.nd receptaculum seminis 
present. Lobes of ovary lon ; ;, flattened, irregular in outline, 
made up of anastomosing tubes. Oo'capt and ootype present. 
Vitellaria loose. . Uterus when ripe with 20-30 lateral pouches on 
either side. 

Uterine pores not seen. 

EGG.-..- With three membranes, outer one -.-45-0.100 mm. 
in diameter, Embryo 0.018-0.021 mm. 


HOST.- llerodia Vhombifer Hollowell, (Type host). 

LOCALITY.- Havana, Illinois. ( Type locality) Type: 

unnumbered "bottles in call, La Rue, Bottle Ho. in Dr. H. 

B. Ward's collection* Slides of same. 

The host a very large female was caught on the hanks of 

at ttxvo-*o- III- 

the Illinois River^and was examined Jul; 9, 1910. From the 

Intestine 57 pienes and entire cestoder were taken . If these 

42 prices had heads attached. ITo strcilas were found with an end- 

proglottid. There were hov/ever a number of specimens with ripe 

proglottids. Because of their translucent appearance in life and 

when preserved in foimol I propose the specific name Ophi o dtaenia 

perspicua^ IIov. Spec. 

The longest specimen of the type of material measured 

36.0 cms. in length when preserved by a max. breadth of 2.0 mm. 

The neck is long, slightly broader than the head and not 

easily distinguished from the segmented portion following. The 

first proglottids are very short. About 16 cms. from the head 


tne proglottids are^ quadrate , length and breadth being about 2.0 
mm. These proglottids are mature. At 26.0 ens. from the head the 
proglottids begin to increase in length and to decrease in breadth. 
The second from the last proglottid measures 3,8 mm. long by 1.2 


mm. broa& k The proglottids here are ripe. 

An examination of other specimens of the lot show a 
considerable variation in dimensions. 


ly Other specimens of this species were furnished me by Mr. 

Hermann Douthitt from IJerodia rhombif er taken in Oklahoma, June 
1910. Two heads of this lot measured respectively 0.374 mm. "broad 
and 0.540 mm. "broad "by 0.27 mm. thick. The suckers measured 0.119 
mm. long "by 0.102 mm. "broad. The neck was 0.277 mm. "broad and 
several millimeters long. The entire length of the longest worm 
was 135 mm, max. breadth 1.190 mm. Specimens were monuted in toto 
and from them a positive identification was made. 


The proglottids are attached by the full width. The 
margins of the strobila are smooth, almost without indentations at 
Lhe junction of the proglottids. As a consequence the segmentation 
is very in istinct, without staining and clearing the proglottid 
limits ^e made out with ifficulty even v/ith a microscope. The 
proglottids are rectangular in shape, rarely oval. The surface 
of the worm preserved is sometimes thrown into shallow longitudinal 
farrows. No transverse furrows are n&ted. 

In life and when preserved in formal the specimens is 
translucent from which character it derives its name. 

The somewhat conical in shape, slightly flatten- 
ed aorsovoutr&lly marked off into four quadrants "by grooves 
which extend from the "basal region to, or nearly' to, the apex. 
Each quadrant hears at its thickest and "broadest region a sucker 
which opens outward and slightly forward. The apex may, does 
not exhibit a circular depression or a fifth sucker. 

The head varies in breadth from 0.255 mm. to 0.408 mm. 
This dimension slightly exceeds the thickness and length. A head 
0.357 broad measured 0.306 mm. in thickness, while a head 0.408 mm. 
broad was about 0.2 . mi . m . 

bo considerable variation in size 
gn shape. In general they are nearly round or Jin outline with 
shallow cavities. However some of them appear nearly triangular. 
The measure ••105-0*170 mm. in diameter. In the smaller ones 
the effect of contraction is very plainly shown by the deeper cavity 


and by the thicknned muscular wall. The diameter of the sucker 
opening varies from 0.053-0.105 mm. 

The neck is always long from 5-7 mm, and its "breadth is 
subject to considerable variation from 0.170-0.425 mm. Since 
this/study is based on toto preparations exclusively the excretory 
nervous and muscular systems were not studied carefully. The 
excretory system is made up of coiling trunks and anastornasing 
brnaches in the head and extreme anterior neck region. In the 
proglottids two pairs of main lateral excretory vessels can be made 

°ut . r , . ~ / 1 

oEXUAl ORGANS.- These organs* in their character and 

relations are typical of the snake Proteocephalids. The genital 

sinus is marginal, irregularly alternating and situated near the 

middle or it^be as far ante-ior as the end of the first 13 of the 
proglottid. It is marked by a Blight elevation which become: more 
pronounced in the more elongated proglottids. The vagina 
and cirrus open into the common shallow genital sinus. The . igina 
lies either anterior or posterior to the cirrus-pouch but more 
frequently anterior. Rarely the vagina is dorsal to the cirrus- 

H ale organs .- These consist of tests, vas 3fferentia, 
Yas deferens, cirrus, and cirrus-pouch. The tests are very 
numerous, 150-215. They are arranged in two fields, leaving 
a free median zone. In elongated mature and ripe proglottids the 
testes are oval or even pclygonal. The measure up to 0.053 x 0.106mm, 


It is to be noted that the tests tend to occur in groups of two 2, 
3, 4, -In these cases the limiting membranes of the testes touch 
each other. Each group is separated by a small interval from the 

The vasa efferentia hich connect the testes with the vas 
deferens may be clearly seen in some toto preparations. They much 
resemble the vasa efferentia In P. Filaroides and other specimens. 

The vas deferens in young proglottids is a tube, nearly 
straight, arising in the middle of the proglottid and extending to 
and intering the cirrus-pouch. As the proglottid becomes older 
this tube becomes longer and more and more coiled until in mature 
and ripe proglottids it forms a heavy mass of coils lying between 
the cirrus -pouch and the middle of the segment. When filled with 
deeply stained spermatozoa, this mass becomes very prominent. Up- 
on entering, the cirrus-pouch the vas deferens is again coiled 
several times before passing over into the he a ior walled cirrus. 

The cirrus is somewhat swollen but is much smaller than 
Gr e p i d o b o t h ium Genardii, P. Calmettei. or P. gr ancles IT 8P. When 
protruded it is slender and of even diameter from tip to base. Its 
length protruded is about 0.200 mm. This measurement is taken from 
a specimen found in Tropidonatu s Rhomb i f or caught in Oklahoma by 
Herman Douthitt and identified by me. 


The cirrus-pouch is more slender than in several other 
Proteoce oh alu s species from snakes. In mature and ripe segments 
it measure from 0.255-0.320 ram. long by Q, 080-0. C J0 mm. hroad. In 
maturing proglottids it measure about 0.250 mm. long and is 
considerably narrower than in ripe segments . Its length goes into 
the proglottid breadth 3-4 times. 

Female organs. - These consist of vagina, a bilobed ovary, 
oocSpt, oviduct, ootypo, shell gjand, uterine passage uterus, 
vitellaria and vitelline ducts. 

The vagina very near it^ opening to the exterior has in 
mature and ripe proglottids a wide but short inflated region. 

V/hether a sphincter nuscle is present has not been determined • 

o f 

In this region the vagina is richly set wit} gland sells. 

In its course to the interovarial space the vagina, bends 
somewhat anteriad and them inward and poster iad crossing the 
numerous coils of the vas deferens but not crossing the cirrus 
pouch. Just before the ovary is eached the vagina is somewhat 
twisted or even thrown into small coils. A small receptaculum 
seminis. can be demonstrated in some preparations. 

The ovarian lobo. are long flattened bodies^ irregular 
utline. The;/ a r e not solid but seem to be made up of short 
heavy anastomosing tubes ox irregular shape. 

Just back of the midpiece of the ovary the muscular 
oorapt can be noted even in toto mounts. The coils and conections 
of the ovicuct, vitelline duct, uterine passage and lower vagina 


cannot be made out distinctly. An ootype and shell gland have been 
seen in some preparations. 

The vitellaria are loosely follicular. The paried 
vitelline ducts can "been seen to cross th ovaries. 

The uterus in maturing proglottids is a median tub' from 

by the method that 
which as the proglottid occomes older lateral pouches arise^ I have 

described for P. filaroidss a drawing (PI. fig. ) shows the method 

of branching of the uterine outpocketing in an early stage. There 


are 20-30 main pouches on either side and a number of smaller ones. 
Ho uterine pores have been seen. In one preparation the uterine 
passage can be traced to a point -.290 mm. anterior to the mid- 
piece of the ovary where it discharges in to the uterus. 

EGGS.- These are furnished with three membranes an outer 
smooth thin anc hyaline, a middle thick and granular and an inner th 
membrane, surrounding the embryo. The outer menbrane measures 0.045 
-0.060-0.100 mm. The embryo 0.018-0.021 mm. in diameter. The outer 

membrane is spheroidal. 

RELaTIOHSHIPS.- This. species is much smaller than 
C r e p i a o b o t x lium G en . rdii CMarenselle ri ; . Calmettei . and 0. 
grandis and 0. Trimeresuri in most repects tho it may bo longer 
than the observed lengths of some of those. It is most nearly like 
0. Mattere^i likewise from one of the Colubrid.-e . It has a larger 
head and larger proglottids, also a larger number of tests which 
are larger than those of the latter. 


. perspicaa is much larger than 0. f ilaroides from 
Aribly stoma t j ftrift um 2 „ird. It has a smaller head and more testes 

than 0_j_ f ilaroides . The position of the gonita??in the two species. 
i,s very different. 

u. ■ perspfcua has a smal er head and ^mailer sucker,* and 

more numerous tes'les than 0^ L onn~ber,g;ii from ITecturus maculatus £af . 


(PI. (o Figs.//yV^3) 

1909: Proteocephalus filaroides La Rue, 1909:17-49. 

SPECIFIC DIAGNOSIS,- Characters of genus, worms attenuate, 
small, thin, flat. Length 80-110 mm. max. breadth about 0.80-0.90 
mm. Color white, somewhat translucent. Strobilation not evident. 
Intersegmental furrows shallow. Surface smooth. Scolex globose, 
flattened dorsoventrally with conical apex, without spical de- 
pression or fifth sucker. No rostellum, no spines. Head not marked 
by furrows. Breadth of head 0.366-0.46 mm. Suckers deep, muscular, 
oval in outline max. diamension 0.165-0.184 mm. Heck narrow 3-4 
mm. long. 

First proglottids 0.30-0.36 mm. broad x 0.10-0.17 mm. 
long. Mature proglottids quadrate or longer than broad. Ripe 
proglottids from 1.6 mm. long x 0.8 mm. broad to 4.0 mm. long x 
0.75 mm. broad. End proglottid present. 

Musculature weak. Parenchyma coarse, filled with large 
fat globules. 

GENITAL ORGANS.- Typical of genus. No genital papilla. 
Genital pore irregularly alternating, situated at end of first fifth 
or proglottid margin. 

Male organs . -Testes 0.05-0.06 mm. in diameter, 70-114 in 
number arranged in two lateral fields. Ductus e jaculatorius with a 
few coils in oirrus-pouch. Cirrus weakly muscled, slender, cylind- 
rcal, 002-0.3 mm. long when protruded. Cirrus-pouch about 0.23 mm. 
long by 0.11 lam. broad. 


Female organs .- Yaginia always anterior to cirrus-pouch, 
not crossing later. Weak sphincter vaginae and small receptaculum 
seminis present. Ovarian lobes thin, alate, composed of anastomos- 
ing tubes. Organs of interovarial space typical of genus. 
Yitellaria with large follicles. Uterus, when developed, with 25- 
35 lateral pouches on either side. Uterine pores 8-12, 

EGGS.- With three membranes, Outer one 0.035-0.100 mm. 
second, 0.030 mm. embryo 0.021 mm. in diameter. 


Habitat.- Intestine and rectun. Ambly stoma tigrinum 
Baird"(type host), Nebraska, (type locality)^ and Kansas. 

TYPE .- Unnumbered aleoholies in coll. La Rue and slide 
or same. Autotype in coil. Dr., H. B. Ward. 

HISTORICAL SUMMARY.- This species was first described 
by La Rue (lyuu:iv-4vj as Proteocepnaips filaroides. So far as I 
aan find out no one else has worked on the species. 

SOURCE OP MATERIAL- This material was obtained by me 1'rom 
Ambly stoma tigrinum Baird caught in ponds in Nebraska, both at 
Crete and in Cherry County^ also in a pond at Belleville, Kansas, 
i These specimens v/ere taken during the summers oi' 1VU7 and. i9U8. 

EXTERNAL CHARACTERS.- 1J These cestodes are aitennate, 
I ther. ana flat. They are white in life at times somewhat translucent 

The strooila presents no evident strobilation. The proglottids 
! are ciosely joined by their full breadth. The intersegmental 
lurrowc are snallow. There ae no longitudinal or tranverse furrows 
! in preserved specimens. 

x'he data presented here was taken almost entirely from 
La Rue ( ly y ) . 

The strobiia is from tfu to liu mm. long by a max breadth 
of u.uu-o.'JU mm. Jne scolex (PI. fig.//jH is gio Dose flattened 
1 dorsoventually with a smooth conical apex„ in which there is no 
depression nor fiitn sucker. There is no rostelium and no spines. 
The surface oi the neaa is usually smooth and but rarely marked by 
furrows. The nead is from mm. broad. 


The i' our suckers are oorne at the broadest part oi the 
scoie-x. They are oval in outline and. they measure irom u.ibt>-o.lH4 
mm. in the maximum dimension. The sucker sai> e deep. Their musculatur 
is well developed. 

As in seven! 3ther species 01 Proteocepnaius which have 
no filth suoker there is a structure (Pl.£> lig.//fr /w ) in the 
tissue beicv; tne surface ol the apex wnich I have lormerly (La Rue 
lyuyj called an end- crgan. Its nature and significance has already 
been discussed in tho general discussion oi the groups (P. ). 

The narrow neck is '6 -co 4 mm. long. This is xoliowed by 
a region oi shoro progiottids which gradually become longer and broad 
The increase in length is more rapid than the increase in breadth* 
nence the progiottids change from "Broader to long than co quadrate 
and -chen to longer xhan broad. The youngest progiottids measure 
o.3u-o.«ib mm. broa^ by 0.10 to 0.17 mm. 

long. Ripe progiottids measure l.b mm. long by u.o mm. broad and 
in some cases as much as 4.8 mm. long by u.Yk mm. broad. An end- 
progiotiid with rounded posterior end may be present. 

INTERNAL ANATOMY.- The musculature of the body is 
weaki.y developed. This is particularly true oi" the longitudjnai 
muscles which lie betv/een the cortical and me dually layers af the 

EXCRETORY SYSTEM.- The excretory system in the head is 
made up oi' coils oi' small anastomosing vessels wnich are connected 
with the two pairs oi mam lateral vessels in the strobila. The 
two pairs oi' lateral vessels lieswithin the dermomuscular sheath. 


The genital canals, cirrus-pouch and vagina always pass between 
the dorsal and ventral vessel, Th> ventral and dorsal vessels 
are connected with the exterior by frequent branches which discharge 
on the dorsal on ventral surface more frequently than on the margin 
of the segment. Regularly situated foramina secudar ja do not 
exist. A transverse excretory commissure in the posterior part of 
the proglottids has been found connecting the ventral vessels. 

Parenchyma .- The parenchyma is coarse. i7ithin the 
parenchyma are many iarge spaces which in material prepared by 
01 dinary methods appear a to be nothing more than abnormally 
iarge parenchyma spaces. V/hen fresh mateiiai is stained with 
"Sudan ill" or treated with osmic acid these spaces are found to 
be fat globules. These tests have likewise been applied by me to 
0. lonnbergii with simuiar resu±ts. I have found the xat globules 
in species of i J roteocephalus but the globules are not so large. 
Testes were also made on Taenia s agr.natea^ Taenia s errata and 
Dipyj-idui.. caninum and fats were found in each case. Testes for 
fat have not been made on the species from snakes but those species 
snov; the loose parenchyma which is one of the characters by which 
the genus is distinguished from Proteoce phaius . 

GENITAL OR GAITS.- These are typical of the genus. There 
is no genital papilla. The genital pore is irregularly alternating. 
It is marginal situated near the end or the anterior fifth oi the 
proglottid. The genital pore leads into a small genital atrium into 
which both cirrus-pouci'. and vagina open. The vagina is always 



anterior zo the cirrus-pouch. 

Male organs. - The testes ( Pl.kf ig./ 22 - J number vu-114. 

They measure about u.Ub-U.Ub mm* in diameter and are situated in two 

broad rields on cither side or a free median zone. The 

testes are dorsal to the uterus. The mass ox" coils of the vas 

deferens extends into the mid-fields of the segment. These coils 


function as a vesieula seminalls. The ductus e Jaculatorius makes 

a few coils before passing over into the straight cirrus. The 

| cirrus is weakly muscled. When protruded it is U.2-U.3 mm. long. 

It is slender and of almost uniform diameter from base to tip. 

The cirrus-pouch is weakly muscled. Its length is about U.22 mm. 

and its breadth about u.n mm. being broadest at its inner end. 

Female organs . - The vagina (PI. io fig./2C) always lies 

anterior to the cirrus-pouch and does not cross tne latter. There 

are no coils of vagina anterior to the inierovarial space. A week 

sphincter vagina and a small receptacuium seminis are present. 

The lobes of the ovary (Pl.£ fig./W) are thin and some- 


wnat alaxe in shape. They are made up of tubules. The 

organs (Pl.£ fig./ ^5) of the interovanal space are -typical of the 
genus. The follicles of the vitellaria are large. 

The uterus is a median tube in mature proglottids. From 
this dtube tnere arise by the process described by la Rue (lyu9: 
33-37) both lateral and ventral diverticula. The lateral diveritcuia 
in fully ripe proglottids extend to the vitellaria which by this 
time nave degenerated to a large extend. They njrmber from zo 3b 


The veircral diverticula number from 8 to 12. These are snorter. 
In time ^hey pierce the ventral body-wal as the uterine pores. 
! The ventral body-wal now gives v/ay along this line oi' peri', rations 
cau. ing a Sift which extends fromm one end oi' the progloxxid to the 
o-cher. Thus the eggs are discharged. 

EGGS.- The eggs are covered with three membranes oi the 
character usual to this group. The outer membranes zr in and 
hyaline varies from 0.036-U.1UU mm. in diameter. 1 1S wells up greatly 
when the eggs is discharged inxo uhe water. The second membrane is 
thick and granular about 0.U30 mm. in diameter. The inner membrane 
is -chin. The six hooked embryo measures abo t u.UHl mm. in diametoi 
DEVELOPMENT . - The larval lorm of this cestode is 
frequently found encysted in the abdominal viscera and body muscles 
oi" the host Amblysxoma tigrinura . They also are found free in the 
body cavity of the same hos"c. They have not yet been round in an 
invertibrare host. 


(PI. f figs./£W*3 ) 

1895: Iohthyotaenia Lonnbergi i Fuhrmann, 1895:218-226. 

1909: Proteocephalus Lbnnborgi i. La Rue, 1900:43. 

SPECIFIC DIAGNOSIS.- Characters of genus. Length up 
to 17.0-19.0 cms. Breadth up to 1.35 mm. Strcbilation usually 
indistinct. Intersegmental furrows shallow. Proglottids attached 
by full width. Sco] ex globose, flattened dor sovent rally, 0.50-0.60 
mm. broad. Ho rostellum, no spines ,no fifth sucker. Suckers 
prominent, muscular, oval or round, deep, 0.24-0.26 mm. long by 
0.14-0.22 mm. broad. Opening 0.20 x 0.10 mm. in oval suckers. 

Heck 0.375-0.54 mm. in minimum breadth', about 2.0 mm. 

long. First proglottids about 0.05 mm. long by 0.50 mm. broad. 

Mature proglottids quadrate, 0.85-1.00 mm. souare, or longer than 

broad measuring as much as 2.5 mm. long by 0.45-0.5 mm. broad. 

Ripe proglottids not seen. 

EXCRETORY SYSTEM.- Two pairs of main lateral vessels*/"^* 
distance from lateral margins 

\of strobila. vessels spiral in form. Transverse commissure present 
in posterior end of each Proglottid. Many branches extending from 

dorsal and ventral vessels to exterior. Anastomosing branches in 

GENITAL 0RGA1IS.- As in genus. Genital pore marginal, 
irregularly alternating, situated at end of first one thirft or 
two fifths of proglottid. 


Male organs .- Testes round or orval, 0.05 -0.08-0.12 
mm. i-n max. dimension. Testes in two fields extending laterad of 
excretory vessels. Midzone of proglottid not entirely free of testes, 
Vas deferens forming a large mass of coils posterior to cirrus- 
pouch. Cirrus-pouch 0.185-0.380 mm. long "by 0.05-0.085-0.10 mm. 
"broad. Ductus ejaculatorius in several coils, frequently forming a 
small vesicula. Cirrus not muscular, cylindrical v/hen protruded, 
0.13-0.15 mm. long x 0.045-0.05 mm. thick. 

Female organs..- Vagina anterior or posterior to cirrus- 
pouch. Vaginal opening sometimes doesal to latter. Vagina never 


crossing cirrus-pouch. A large dilation in vagina of some proglottid', 
Sp'nincter vaginae and receptaculum seminis present. 

Ovary thin, flat, lobes wedge-shaped or alate. Organs 
of interovarial space ty ^ical of genus. Vitellaria sparse. Uterus 
with 25-40 lateral pouches. 

EGGS.- Yol : naas 0.010-0.012 mm. in diameter. Egg 
membrane s could not be seen. ITo embryo seen. 

HABITAT.- In intestine of ITecturus maculatus aaf . 
Fuhrmann does not state the locality of his type specimens. I have 
frequently found the species in Necturi from Ohio and Indiana brought 
to the laboratory for dissectior. Fuhrmann' s specimen was obtained 
from Prof. F. Zschokke who -received it from Prof. R. Burckhardt. 

TYPE.- Prepared slides in collection of Fuhrmann. 


HISTORICAL SUMMARY • - Fuhrmann (1895-218-226) described 
and delineated this species, calling it I chthyo taeni a Lonnbergii. 
It has not been redescribed until the present. 

SOURCE OP MATERIAL.- This fctudy is based upon alcoholics 
and prepared slides in coll. of La Rue. This material has been care- 
fully compared with Fuhrmann 's preparations oi" the type «hich Prof. 
V/ard very kindly secured for my use. I have been able thus to 

identify positively my material as being identical with Fuhrmann's. 
Reference will be made from time to time to the work of Fuhrmann 
(1895) from which certain data were secured. 

EXTERNAL CHARACTERS.- This form is much more robust 
than 0. f ilaroides . Its length may be as much as 17.0 cms. or more. 
Fuhrmann f s specimen which was immature measured 19.0 cms. The 
worm is thin and flat. Its breadth varies up to 1.275 mm ., Fuhrmann 1 s 
material had a max. breadth of 1.35 mm. The strobilation is usually 
not distinct. The margins of the strobila are smooth. Rarely the 
posterior part of the worm maj show a distinct segmentation. The 
proglottids are attached by their full width. There are very few 
transverse folds but shallow longitudinal furrows are not uncommon. 

The scolex^is somewhat globose and is flattened dorso- 
ventrally. The suckers are prominent and situated at the broadest 
part of the head. The breadth of the scoley varies from 0.60- 
0.60 mm. There is no rostellum, no spines > and no fifth sucker. The 
suckers are round or o . al in outline. The cavity is deep, the 
musculature strong. The length of the suckers varies from. 0.24-0.26. 


mm. and the breadth from 0.14 to 0.22 mm. The opening measures 
about 0.20 x 0.10 mm. in the more oval suckers. 

The neck immediately behind the head is from 0.375 to 
□•54 mm. "broad. Its length to the first traces of segmentation is 
about 2.0 mm. 

The young proglottids measure 0.27 mm. in length according 
to Fuhrmann. I found that the first discernible proglcttids are about 
0.050 mm. long by 0.50 mm. broad. The length and breadth of the 
proglottids increase rapidly for a distance , then the length increase 
and the breadth decreases. In my material the max* breadth of 1.275 
mm. was reached long before the proglottids were mature. Proglcttids 
in this region measure 1.275 mm. broad by 0.30-0.40 mm. long. 
Mature proglottids are about 0.85-1.0 mm. square or they may be 
longer than bread. Some of the elongated proglottid' measure 2.5 
mm. long by 0.45 to 0.5 mm. broad. In Fuhrmann's specimens the last 
proglottids were 0.7 mm. long by 1.35 ram. broad. Neither Puhrmann 
'not I have seen any ripe proglottids. All the hosts examined by me 


i had been caught for some time. They were examined in the spring 
months. It may be that they attain the sexually ripe condition later 
in the season. 

pSTSR&KL AH ATOM!.- Excretory system . The excretory 
system is made up of two pairs i" main lateral vessels, dorsal and 
ventral. These are situated much farther mesad than is usually the 
case in this group. These vessels pass thru the tissue in the form 
of loose spirals. 


! Prom them at frequent intervals small branches pass to the dorsal 
and ventral surfaces. A transverse commissure is present in the 
posterior end of 3ach proglottid. In the head the main excretory 
vessels are connected with many anastomosing tranches. 

Parenchyma and musculature ; The parenchyma resembles that 
of _0. f ilaroides . Chalkbodies of lar&e size are abundant especially 
in the head and neck. The musculature is weak. 

GE1TITA1 ORGANS.- The genital pore is irregularly 
alternating, is marginal, and is situated at the end of the first 
third or two-fifth of the proglottid. The cirrus-pouch and vagina 
pass to the exterior between the ventral and the dorsal excretory 

Male organs.- The male organs mature first. The 
testes^ number from about 90 o 150>the number varying greatly in th 
proglottids of the same strobila. They measure from 0.05 to 0.080 
and even up to 0.120 mm. in maximum dimension in elongated testes. 
The testes lie in two broad fields* Each field extends laterad and 
i meaad of the excretor vessels. In many proglottids an occasional 
testis lies in the midfield. The testes are dorsal to the uterus. 

The vas deferens in mature proglottids forms a large 
mass of coils which function as a vesicula seminis. This mass 
of coils is usually situated posterior to the cirrus-poach • It lies 
almost in the midfield. 


The cirrus pouch is short and "broad. Its length varies 
from 0.185-0,280 mm. and its breadth from 0.050 mm. in the case of 
the longest pouches to O.0C5-0.100 mm. in the shorter pouches. The 
cirrus-sheath is broadest at the inner end. 

The ductus ejaculatorius forms several coils and then it 
passes over into the slender cirrus. In some proglottids the 
ductus was dilated to form a small vesicle. 

The cirrus itself is not very muscular. When protruded 
it is cylindrical and ahout 0.13-0.15 mm. long by 0.45-0.05 mm. thick 

Femal e organs t- The vagina^lies anterior or posterior 
to the cirrus-pouch. Its opening is usually dorsal to the pouch. 
When the vagina lies anterior to the cirrus-pouch it does not cross 
the latter. Its passes below the coils of vas deferens. In many 
mature proglottids the beginning part of the vagina is greatly 
j dilated and sometimes This dilatation extends for some distance 
dorm the length of the vagina. There are no coils of the vagina 
j anterior to the ovary but that duct may lie in a series of sinuous 
curves. A weak sphincter vanina close to the external opening and 
a small receptaculum seminis near the mid-piece of the ouary are 

The shape of the ovary varies with the state of contraction 
of the proglottid. J-t is always ztiln and flat and ir made up of 
anastomosing tubules as in 0. filaroides . In very "broad but short 
proglottids the lobes of the ovary are long, slender , wedge-shaped 
ffith the apices in the mid-field. 


In attenuated proglottids the lobes are much shortened and "broader. 
In this case they may be distinctly alate. 

The organs of the interovarial space are as delineated for 
0. filaroides (PI. 6> fig. /2-3 )• The vitellaria are sparse. 
The follicles are of medium size. 

The uteruo has not been observed in ripe proglottids. 
In one proglottid in which a few eggs had been passed into the 
uterus there were from 25 to 40 lateral uterine pouches on either 
side. Ho uterine pores have been observed. 

EGGS.- Ho eggs with developed embryos could be found, 
A few eggs which could be but poorly seen within the uterus showed 
yolk masses v/hich measured from 0.010-0.012 mm. in diameter. The 
egg membrane could not be measured, Fuhrmann saw no eggs. 

RELATIONSHIPS: - This species tho most closely related 
to 0. filaroides can be readily distinguished from the latter by 
its large strobila, large head, larger suckers, and larger testes. 
The excretory vessels are spirals in 0. Lonnbergii but straight in 
0. Filayoidfc ?, The shape of the ovarian lobules is also different. 
0. Lonnberg-ii differs from the Ophiotaenia from snakes by the posi- 
tion of its genital pore, and by the less muscular character of its 
cirrus. Its ovary is also of a different shape. The position of 
the excretory ducts is much farther mesad in this species than in 


any of the species from snakes known at present. 

Further investigation may prove that the proteocephalids 
infesting the Amphibia do not belonp; with the Ophiotaenia . The 
extreme anterior position of the genital pore and the cylindrical 


and weakly muscled cirrus in 0. f ilaro ides . and 0. Lbnnbergii 

makes" these species different from the species from snakes. In 

the latter species the genital pore is near the middle and the. 

cirrus is either thick and muscular or slender with a muscular and 

bulbous base. At present I do not feel justified in erecting a 

new genua for the^ f i oteocephalids a&d since their closest allies are 
the Ophiotaenia I shall place them in that genus. 


(PI. (o 21gwf,M-/*t) 

SPECIFIC DIAGNOSIS.- Characters of genus. Body very 

long (fragments 200 mm.) by 2.75-4.25 mm. broad in region of ripe 

proglottids. Heck 5-8 mm. long. First proglottids much broader than 

long; proglottids with developing sex organs quadrate or nearly so; 

ripe proglottids quadrate to mud": longer than broad. Head large 1.0- 

1.2 mm. broad at base of suckers. No fifth sucker or rostellum, no 

hooks. Suckers nearly circular, deep, and muscular, 0.340x0. 3 60 mm. 

Genital aperture marginal, irregularil; alternating, situated very 

near middle of proglottid. Ho genital papilla but sometimes a 

deep depression • Testes large, numerous, 200-250 in number. Arranged 

in tv/o broad lateral fields. Cirrus-pouch 0.24-0.26 run. broad 0.64- 

0.75 mm. long length equal to l/3-l '5 proglottid width. Cirrus 

short and heavy. Few or no coils of va^ deferens i] cirrus-pouch. 


Vagina anterior^ posterior to cirrus-pouch", Sphincter vaginae heavy. 
Uterus with 40-60 lateral out pocket ings on either side :. ventral 
uterine openings 2-8. 

EGOS.-Out'jr membrane ellipsoidal 0.026-0.037 mrn.or some- 
times spherical 0.032 mm. in diameter; second membrane CO:, 1-0. 023 

mm., Embryo 0.015-0.016 mm. Outer membrane smooth. Type. U. S. 

B.A.I. 14854. 

Habitat.- In intestine of Anciatrouon piscivorous, (type 
host) Nat. Zoo. park.- The locality from which this host came 
originally is not known- 



The species is found onl; in North America. 

SOURCE OF MATERIAL.- Six or eight heads together with 

numerous pieces in bottle be. ring U.S. B.A.I, number 14854 were 

labelled Iohthyotaenia : host ^npis trodon piscivorous ; location, 

intestine; locality Nat. Zoo Park, collected by 3. S. Shauhan, 

iTov. 18, 1907. These v/ere secured lor my study by Dr. Henry B. Ward. 

EXTERNAL CHARACTERS*- No complete worm was present, 
The largest piece was 2,00 long ^nd 

3,5mm. broad at the anterior a by 4.25 mm. broad at the posterior 
end. It was made up of nearly mature, mature and nearly ripe 
proglottids. The strobilation was not evident at the anterior end, 
but clear after 60 C»r 70 mm. Prom the size of the proglottids and 
their state of development it is likely that the whole worm wotild 
attain a length of 300-400 mm., perhaps more. 

Like other Proteocophalus species inhabiting snakes and 
amphibia the body is quite flattened. In this case the body is 
thicker than I have observed in P .racemosa ,Ge rrardii m p . gattera 
P. Tr: i moresuii , P. perspicuus. 

The heed is large tho not so large as in 0. Harenzollor i 
(Baarois) or Crgjoidobothr iu a Gg rrardii (Baird), It measures 1.0-1.0 
mm. in diameter at the base of the euckers. The length of the 
scMex from the tip to the base of the suckers measures 0.50-0.60 
mm. The -e is no rostellum,no hooks, no fifth sucker, and from 
toto mounts an endorgan was not observed. No sections have been 
made . 


The four deep suckers a e borne on the "broadest part of the head 
and these are directed outward. They are not prominent. They 

I are strongly muscular, nearly circular in outline and without the 
upward pointing projection that is found in C. Gerrardii . The 
suckers measure 0.34-0.35 mm. in transverse diameter "by 0.35-0.36 
mm. in longitudinal diameter. The sucker cavity is deep. Its 
opening is 0.15-0.17 mm. in longitudinal diameter by 0.20-0.22 mm. 
in transverse diameter. 

Immediately back of the he<. d there is sometimes a large 

. inflation. Since this is not always present it must be due to 
contraction of the heavy longitudinal mu: alec of the upper part of 
the neck. The neck itself "is broad, thin and long being 0.85-1.0 
ram. broad by 4-8 mm. long. 

The first proglottids are quite short being several times 
broader than long. This condition soon changes, the proglottids 

! become nearly quadrate or quadrate. Nearly mature or mature and 
ripe proglottids are longer than broad. Those proglottids in 
which genitalia, ( Vagina and cirrus-pouch) begin to show measure 1 mm. 
broad by 0.5 mm. long. Those proglottids in which the anlage of the 
tests may be seen measure 0.850 mm. broad by 0.680 mm. long. A 
few nearly r$.pe proglottids are broader than long, measuring 3.25 
mm. long by 4.25 mm. broad. The longest proglottid measured 8 mm. 

long by 2,75 mm. broad. Many proglottids measure 3-5 ram. long, by 
2-3 mm. broad. 


The surface of the "body is somewhat wrinkled, lae perhaps 
to being twisted for a long time. There seem normally to "bo no 
furrows > either longitudinal or transverse. The proglottid limits 
are not plain in young proglottid^ hut quite plain in the older ones. 

The genital aperture is marginal, irregularly alternating 
from right to left. Frequently the genital aperture appears on 

the same side in several suc-es^ive proglottids. As many as ten 


successive segments have "been .vith the genital aperture 

on the left side. This side seems to he the one favored for the 
situation of the genital opening. The genital aperture is not 
marked by a papilla. Indeed, in many proglottids the muscles are 
so contracted about the genital aperture as t - cause a deep 
depression (PI. fig* )• In such cases it is to be noted that the 
cirrus and vagina open to the exterior separately. In any case it 
is doubtful if a true genital cinus exists. If it exists it is 
extremely shallow. 

IHTERHAL AH ATOMY.- Genital o rgans ; The arrangement of 
the sexual organs agrees in general with that of other Proteocephalid 

Male organ s.- The cirrus-pouch^is short and broad being 
broader at the distal end. It is either straight or curved de- 
fend"' ng upon the state of contraction of the proglottid and of the 
muscles surrounding the genital aperture. When a depression about 
the genital aperture exists the cirrus-pouch is usua'ly somewhat 
bent. It measure from 0.64-0.75 mm. long by 0.24-0.2G mm. broad. 


The natio of its length to the breadth of the proglottid is 

The cirrus ( Pl.£ f ig. / 3 ) itself when it is protrufced is 
thick, short, straight^ and heavily muscled. It is continuous with 
the ductus e jaculatorius which forms no^ or at least few, coils 
within the cirrus pouch. The drawings (PI. (e figs./?*,/*) show only 
a slight twist or turn in this part of vas deferens. These 
features show the cirrus and vas deferens to he very different 
irom these organs in 0. ■■■Iarenzelleri w ith which this species is 
closely allied. In thai; species there are many coils of vas 
deferens within. the cirrus-pouch. The cirrus when protruded is 
thick at the base, drawn out to a long slender tip. Schwarz errs 
in saying that the cirrus-pouch is evaginated. His own figures 
Show the cirrus-pouch in situ. Outside the cirrus-pouch the vas 
deferens is thrown in", o mnerous coils which serve as a vesicula 

The testes which lie in two fields are very numerous, 
numbering 200-250. Thqpmeasure 0.04-0.05 mm. "broad x 0.075-0.100- 
8.125 mm. long. 

Female organs .- The vitellaria are follicular in 
structure, the follicles not being grouped in any special manner. 
Each follicle measures 0.015x 0.030-0.035 mm. 

The vagina (PI. & f ig$./2£/?/) opens to the exterior 
either anterior or posterior to the cirrus-pouch. Out of 45 


proglottids the vagina in 21 casecfopened anterior and in 24 
cases- posterior to the cirrus-pouch. A heavy sphincter vaginae 
0,095 mm. long x 0.150-0.160 broad (/agina include dj surrounds 
the vagina near opening to the exterior. The sphincter itself 
is from 0.040-0.080 mm. thick. 

A diagram A f rom A tcCto preparation shows the arrangements 
of the organs oi the intorovarial space to he similar to that 
figured for other Proteocephalial species. The ovaries are thin 
dorsoventially. The mid-piece is slender and long. 

The utsrus A in maturing proglottids is a median ventral 
tube. The uterine outpocket ings are numerous there being 40-60 
on either side. They are formed as I have described (La ;.ue 1909) 
foi 0. f ilaroides . No eggs are passed into the uterus until the 
outpocketings have attained LOuie.size. Two or more preformed 
ventral openings to the exterior are developed before many eg-gs 
are present in the uterus • In one ripe proglottid studied there 
were eight uterine openings. 

EGG3.- The eggs taken from the uterus are usually 
ellipsoidal measuring about 0.026x0.037 mm. over the outer membrane, 
Spherical egg! measure about 0.032 mm. The second membrane is 
spherical, .021-0.023 mm. in diameter. The embryo is 0.015-0.016 
in diameter. The mature egs have smooth contours, there being 

no booklets or holding organs as Schwarz (1003) figures for the eggs 

of 0^ Hatter eri (Parojia) 


RELATIONSHIPS . - 0. grandis differs from Q.Marenzelie rii 
(3£-nT>ib) in having a smaller heed, smaller suckers, larger 
proglottids, greater breadth, more numerous tecles, raore numerous 
uterine pouches. The character of the cirrus in the two species 
when evap-inated in very different* 

0. grandis differs from Cropidobothiu rn gerr ardii in 
head and the size of the suckers, and in the character of the 
latter. dgrandis has a long unsegmented neck while 0. Genardii 
has almost no neck. 

0. grandis somewhat resembles 0. Calmet Lei in the size 
of its head and suckers. It differs from that species in the 
size of strobila and of proglotlrids and in length ©f neck, 0. 
grandi s has more tests and a cirrus of different character. 

0. grandis is t much larger than 0. perspacua n. Sp., 
0. Ilatter eri(Parona) 0. t rira^resuri (Parona) and Q. raceme sus 
(Rud.) that a possiblity of confusing it with any of those species 
is precluded. 



(Pi. % Vlsa./fts/fi) 

18V3: Iohthyotaenia Marenzelleri Barrois, 18'Jtf:2-3 

l l J08: Iohthyotaenia Marenzelleri Schwarz, l ( 'UJ:2b-2V. 

SPECIFIC DIAGUOSIS.- Characters of genus. Observed 

j length oi strobila up to 4U cms. Max. breadth 1.5-2.-U ram. Scoiex 

large, round, muscular, 1.0-2. o mm. broad. Well set off irora neck, 

No rosteilum^no hooks. Suckeis four, 0.6U-0 7U mm. in diameter, 
neck, no data. 


Strobiiation distinct. Proglottids closely attched. Mas . 
length or progiottids 0.5 mm. max. breadtn of same 1 .5-2.0 mm. 

Excretory system made up of two parts of lateral vessels. 
GENITAL ORGANS!- Genital aperture irregularly alternate i 
situated near middle of lateral margin of proglottid. 

Male organs .- Tos-es in two lateral fields, 150-200-240 
in number, u.u6-u.uv mm. in diameter. Cirrus-pouch large, extend- 
ing about i/3 across roglottid breadth. Cirru^ large at 
base, filiform at ti. , i mm. long. 

Female orgt as : - Vagina usually posterior to cirrus- 
pouch. Uterus wnen completely developed with 20-25 lateral out- 
pocketing, on either side. 

EGC-S.- With two membranes. Diameter not known. 
HABITAT.- In intestine of Ancistrodon piciscivorous 
Holbr.. Southern United States. 


HISTORICAL SUMMARY.- Barrois U8U8) in a very brief 

description unaccompanied by drawings proposed this species. 

Schwarz (iyu8: 26-27) redescribed this species from Barrois material 

and his description was accompanied by three figures. 

SOURCE OF MATERIAL.- The material which Barrois and 

Schwarz had for study was collected by Doctor Caimette, Dec. .2, 

±8"J7 from Ancistrodon piscivorous HuxDr, a snake indigenous to 

Southern United States. 

Unfortunately specimens of this species could not be 

had by me for study hence the data used in the following description 

are derived only from the articles of Barrois (18U8) and Schwarz 


(iyu8). A stjtdy oi new material is desir A ble in order that a more 
complete knowledge of this i'orm may be had. 

EXTERNAL CHARACTERS.- This is one of the largest of the 
species oi Ojohi_pjLa£iiia. The observed length is about 4u cms. 
Schwarz (iy(J8:26J thought that perhaps the length might exce ed that 
of 0. Coimettei ( 35-8U cms.) The maximum breadth observed was 
1.2-2.0 mm. The scot ex is large, round and muscular. It has a 
bceadth of 1.5-3,0 mm. Barrois described an unarmed rostellum. 
Schwarz found no rostellum, nor does his delineation oi the 
head, which is reproduced. (PI. V iig./ff), show one. 

The scolex is clearly set off from the neck. The 
suckers, four in number, are muscular, prominent and nearly circular. 


They measure 0.60-0. VO mm. No fifth sucker is present. 

The neck v/as not described by Barrois or by Schwarz. 

The strobilation is distinct. The strobila has no 
indentations. The proglottids are strongly attached. The length 
of the largest proglottids is about 5.5 mm. and the max. breadth is 
about l.f-S.O mm. 

The excretory system is made up of two pairs of lateral 

vessels, a dorsal and a ventral. 


3?he genital aperture alternates irregularly. It is 

situated about the middle of the proglottid margin. 

Male organs .- The testes A lie in twp well marked lateral 

fields between the vitellaria and the uterus. Barrois gives 230- 

240 as their number, Schwarz 15( -200. They measure 0.060-0.070 mm. 

Of the cirrus-sneatJ: and vas deferens Schwarz says 

"the relation of the cirrus-sheath and vas deferens is especially 

characteristic for this form. In ripe segments the gre .ter part 

of the cirrus-sheath is completely evaginated. There seeems 

to exist a special miscular apparatus which daaws the cirrus-sheatl: 

when completely sexualy mature from the proglottid, so that it 

projects free from the margin. In young segments it lies in its 

normal position within the segment". 

Schv/arz has evidently misinterpreted the appearance 

in this case. His own drawings show no evigination of the cirrus- 
ouch but a complete evagination of the cir -us which is thick at 
the base, and filiform at the tip. 


Prom this heavy base the ductus ejaculatorius runs straight thru 
the now greatly shrunken cirrus-pouch. There are some coil: of the 
ductus ejaculatorus within the basal part of the cirrus itself. 
I have observed this condition in 0. t rimeresuri 

Schwarz states tha: the cirrus is ab ut 1 mm* long in 
<J 2- m renzeller i. In Schwarz' s drawing the cirrus-pouch extends about 
1/3 across the proglottid breadth. The vas deferens forms a mass 
of coils before entering the cirrus -pouch. 

Fenale organs .- The vagina^ open usually posterior to the 
cirrus-sheath . Its course to the middle of the proglottid 
forms an arc. Az the middle of the proglottid it bends sharply 
posteriad and then its course is direct to the interovarialspace. 

The uterus is a median tube from which arise on either 
side many lateral pouches; 20-2i>, are figured in Schwarz* s drawing 

The ovary is relatively larger. The lobes are 
somewhat winglike in shape. 

EGO-S.- The eggs have two membranes. Ho processes 
arise from the outer one. Ho measurements of the eggs were 
giren by Barrois or by Schwarz. 

RELATIONSHIPS.- This species is clearly differentiated 

from the other large species of the genus by the size and chape of 

head and suckers. The relations of the cirrus-pouch and cirrus 

in other large species of the 
are greatly different genus • £• grandir — -e^bles it in having 


in having the same host species and in being very large. 0. 
grandis has however a very much smaller head which is not readily 
distinguished from the strohila. It has much smaller suckers^ and 
greatly different relations of cirrus and cirrus-pouch. Its cirrus 
as short and thick. It also has very many more uteriae out pocket » 

0. Marenzelleri is not to "be mistaken for 0. r acemosa 
0. N attereri . 0. pigmenta l a 0. perspicua on account of its much 
larger size. 0. filaroiG .es and 0. Lonnbergii being from am- 
phibia and also being much smaller cannot be confused v/ith 0. 
Marenzelleri A 


(PI. % ?igs./57 7 /?t J /6f) 
1901: Ichthyc t aenia Natt ereri Parona, 1901: 4 - 6 . 

1906: Ichthycta enia Batterer! Schwarz, I9O8: 17-19. 

SPECIFIC DIAGNOSIS.- Characters cf genus. Length 75-250 
mm, max. breadth not over 1 mm. Scolex unarmed, rounded but not 
spheroidal, not easily distinguished from the neck. Diameter of 
scolex 0.240-0.250 mm. Suckers, four in number, circular^ cavity 
deep, margin thick, diameter of suckers 0.120-0.150 mm. Ho 
rostellum and no fifth sucker. Heck long, 7-8 times the length 
of the head. 

First segments broader than long, more or less indistinct. 
Mature prcglottids nearly quadrat e- Ripe proglottids longer tnan 
broad, length of so ue up to 2.0 mm. by a breadth of 0.33-0.50 mm. 
Last proglottids readily detached. 

GENITAL ORGANS.- As in genus. Genital aperture irregular 
ly alternating, situated somwhat anterior to middle of proglottid 

Mgl e organs .- Testes 80-100, 0.042 mm. in diameter, 
situated in two lateral fields. Ves deferens a thick mass of coils 
extending even past the middle of the proglottid. Ductus 
e jaculat orius much coiled. Protruded cirrus slender, heavy at 
base, about 0.20 mm. long. Cirrus-pcuch about 0.265-0.280 mm. 
long, extending from l/3 - 2/7 across proglottid breadth. 

Female organs.- Vagina anterior or posterior to cirrus 
pouch. Sphincter vaginae present. Uterus, when fully developed, 
with 15-20 lateral pouches on either side. 

EGGS.- With three membranes. Outer membrane covered 
with fine hooklets. Eggs 0.024 mm. in diameter. 

HABIT/T.- In intestine of Coluber sp from Liguria, a 
district of Italy. The material was collected by Parona in De- 
cember, 1897. 

HISTORICAL SIJMMARY.- This species was first described 
by porona (1901:4 - 6). The description was not accompanied by 
drawings. Schwarz (1908: 17-19) delineated and redescribed the 
species, using Parona* s specimens. 

SOURCE 0? DATA. - The data upon which this description 
is based were secured from the descriptions of Farona (1901) and 
Schwarz (1908) , and from observations made upon slides prepared 
from type material which Prof. H. B. Ward secured from Professor 
Parona . 

EXTERNAL CHARACTERS . - This- cestode is thin and slender. 
Its observed length varies from 75-250 mm. and its max. breadth 
is not over 1 mm. The segmentation is distinct in the posterior 
region. The single proglottids are thin, and somewhat translucent. 
The first proglottids are broader than long and their boundaries 
between segments are not distinct. More mature proglottids are 
nearly quadrate while ripe proglottids are longer than broad. The 
length of ripe proglottids may be as much as 2.C mm. and the 


breadth 0.33-0.50 mm. The last proglcttids are easily detached 
from the chain./ „ i 

The head^is smalll and not clearly £>et off from the neck. 
It is unarmed, rounded scmewnat but not spherical, its diameter is 
0.24-0.25 mm. It bears four circula suckers which have a deep 
cavity and a thick muscular wall. The diameter of the suckers 
varies from 0.120-0.150 mm. There is no rostellum and no fifth 

The neck is seven or eight times as long as the head. 

Cuticula, musclature, nervous system, and excretory 
system are very similar in arrangement and character to the same 
systems in other species of the genus. 

GEFITAL ORGANS.- These in their general relations are 

t3'-pical of the genus. The early developing sex organs may- be seen 

in quadrate proglcttids. The genital aperture lies somewhat anter- 


icr to the middle of the margin of \ segment and it irregularly al- 
ternates from right to left. 

The testes lie in two fields within the vitellaria. The 
median zone of the proglottid is left free of them. They number 
80-100 and measure 0.042 mm. 

The vas deferens forms a thick heavy coil extending from 

the cirrus-pouch even past the middle of the proglottid Within the 
cirrus pouch there is a very connlicated 

mass cf coils of ductus ejaculat bvftus . This is much mere highly 
developed than- in any other species of Ophiotaenia . The cirrus 
when protruded is heavy at the base, more slender at the tip and 


about 0.200 - 0.210 mm. long. In my preparations it measured 0.108 
mm long. It was incompletely protruded. 

and oblong, proglottids its length goes into the proglottid breadth 
from 3 to 3.5 times. / 

pore. It lies either anterior or posterior to the cirrus-pouch. 
In 12 out of 18 proglottids examined by me the vagina had a pos- 
terior position. 

At the distal end of the vagina a sphincter of good size 
is present. The course of the vagina is nearly direct to the middle 
of the segment frequently crossing the interior end of the cirrus- 
pouch or even ljang above the cirrus pouch for its full length. 
From the middle of the segment its course is directly posteriad 
with sometimes a single loose ceil just above or anterior to the 
ovaries. In the int erovaria-,1 space are the usual coils of the 

va-ina, oviduct , uterine passage, and unpaired vitelline duct; here 

The uterus appears in mature proglottids as a median tube 
from which later 15-20 lateral pouches arise on either side. 

The ovarian lotas are a fairly dense compact mas? . In 
many proglottids the ovary has a more elongated and pointed form 
than Schwarz delineates it. 

The vitellaria are lateral, follicular masses, reaching 
from the anterior end nearly to the posterior end of the segment . 

The cirrus-pcuch is 0.265-0.280 mm. long. In quadrate 

also are the oocapt, ootype, and shell-gland. 


EGGS,- According to Schwarz the egg have three membranes 
The diameter of the egg is 0.024 mm. The outer membrane- of ripe 
eggs is covered with fine hocklets or processes which terminate in 
small knobs. I was unable to observe these processes in my toto 
mount s . 

This species is easily differentiated from Crepidobpthrium 

Gerrardi i , 0. Marer zell er i , 0. Ca lmet t ei , 0. rac eme sa , 0. 
Trimeresura and 0. grardis , by its much smaller size and from 0. 
perspicua by its slightly smaller head and suckers, by its lesser 
number of smaller testes, its more voluminous cirrus-pcuch, and 
lesser number of uterine pouches. In many respects the last two 
are quite similar. . Hatt ereri has a smaller head and smaller 
suckers than 0. pigment at a . Further comparisons with O. pigmentata 
can not be made because of the incompleteness of the data on the 



(pi. 7 ti&«/3i'inp/.*>f<?'t* f ~ fi * J 

lbyu: I chthyo taenia Calmettei Barrois , 1-3. 

18ya: I out hyo taeni a Raillfeiti Marotel, 18y8: 99-101. 

18V8: Ichthyotaenia Calmettei Luhe I8y8:652. 

l«y«: Ichthyotaenia Calmettei Marotel, 18yy : 34-42 • 

lyui: Iohthyotaenia raoemosa Parona, iyoi:3. 

iy08: Ichthyotaenia Calmettei Schwarz, iy08: 24-26. 

SPECIFIC DIAGNOSIS.- Characters of genus. Observed length 
0.97-1.2-2.0 mm. progiottids numerous, first broader 

from 27-4U and even up to 8u cms. Max. breadth^ than long, subsequent 
ones quadrate or even much longer than broad. Mature progiottids 
about U.85 mm. square. Ripe progiottids 2-3-4 mm. long by 1.0-1.2 
mm. broad. Strobilation not clear. 

Scolex spheroidal or tetragonal, without rostellum, with- 
out hooks, without fifth sucker. Anterior face of same flattened 
with small elevation at center. Head 1.0-1.3 mm. broad u.t>u mm. 
long, 0.v3b mm. thick. Suckers four, somewhat globular, with deep 
cavity. Diameter of sucker opening 0.12-0.17-0.185 mm. Diameter 
of sucker 0.27-0. 300-0. 408 mm. Neck 0.580-0. J80 mm. broad .by 4-5-8 
mm. long. 

GEHITAL ORGANS.- Typical of genus. Genital aperture 
irregularly alternating, situated at or near middle of proglottid 
margin. Genital sinus shallow. Vagina anterior or posterior to 
cirrus-pouch. Vaginal opening frequently dorsal to cirrus-pouch. 


Male organs .- Testes 130-lbU in number, arranged in tv/o 
lateral fields. Size or testes 0.026-0.04-0.06 x 0.053-0.07 mm. 
Vas deferens a mass of coils extending from cirrus-pouch to mid- 
field. Cirrus-poucn 0.25-0. 29-0 .33 mm. long by 0.100-0.136 mm. 

broad. Ratio of length of cirrus to proglottid breadth from lt6-l:5 

ids to 

in mature proglott^l:4 in ripe proglottids. Ductus ejaculatorius 
with numerous coils. Cirrus broadjmuscular , not seen protruded. 

Female organs. - Vagina broad at distal end, never 
crossing cirrus-pouch in course to mid-field, Course of vagina 
sinuous to interovarial space. Sphincter vaginae present. 
Recaptaculum ceminis not seen. Ovarian lobes slender in young 
proglottids, toroad in old. Viteliaria with small follicles. 

Uterus when fully developed with 24-35 lateral pouches 
on either side. Uterine pores ventral, 2-4-5- in number. 

EGGS.- With three membranes. Outer membrane variable in 
size, second one thicker, 0.022-0.024 mm. in diameter, inner one 
granular^, irreguiar,,0.ul4 mm. 9 embryo U.U12 mm. 

HABITAT: In intestine of Bothrops lanceolatus L. 





Bothrops lance oi atusL 


S. Paulo 


Adolf Lutz 

La Rue. 


HISTORICAL SUMMARY. - Barrois { 18y8 J description or 
this species while meagre and unaccor-panied by drawings agrees 
fairly .ell with the description oi' later invetigators . His 
specimens came i'rom Bothrops lanceolate s L. from Martinique i"rom 
which tiosi; they were secured by Prol'essor Calmette. 

Later in the same year Marotel (18y8j briefly described 
this species under the name I* Raillieti . His specimens came from 
Bothrops lanceoiatus L. from Martin:' que. Luhe ^18V8J called attention 
to the fact that I. Raiileti was a synonym oi" I. Calmettei and 
he included the form in his list of species of Icht h yotaenia . 
Marotel ( 18y<9J34-42j in a mor>. extended paper described this species. 
His paper is accompanied by several drawings. 

Parona (J 9J1J listed this species in a record of parasites 
from South America under the name of Ichthyotaenia racemosa . His 
specimens are from B. lanceoiatus from . S. Pauilc, Brazil. 
Schwarz ( lyua : 24-2b] added but littie to the work of Barrois and 
Marotel. He determined that there was no rostellum. 

SOURCE OF MATERIAL.- This study is based on material 
which Prof. H. B. Ward secured for my study from Professor Parona. 
It bore the label. n T.( Oocnoristica ) racemosa 

S. Paulo, race. A. Lutz." This is evidently the material 
on which Parona (lyui) based his report. There he stated that it 
came from Bothrops lanceaiatus L. The material consisted of seven 
or eight pieces to one or wnncn a head was attached. The head was 
cleared in glycerine and studied in that condition and some or the 
smaller pieces were stained and mounted as toto preparations. 


The identification or tne manorial was made from These preparations. 
The report is also Dased upon data secured from Barrois (luyu)^ 
Marotel dayy) and Sohwarz (lyoa). 

EXTERNAL CHARACTERS.- 'l'he longest piece which I observed 
measured £70 mm. long by 2.u mm. broad. Barrois (18yd ) states that 
he observed a iengtn or 35-tiU centimeters by a breadth or O.y/ mm. 
tor this species, Marotel 1 s( layy ) specimens measured up to 35-4U cms. 
long by a max. breadth or 1.2 mm. Schwarz (lyoa} reported a breadth 
of 1.5 mm. 

The strooila is made up or many progiottids^ as many as 
Htjy ( Barrois j. Tne progiottids are closely joined to one anotner. 
As a rule no inter-segmentai furrows can be seen with the unaided 
eye. In a single piece the strobila was strongly contracted and 
nere tne strobiiation was evident. The suriace or many of the 
progiottids is tnrown into numerous longitudinal loids. 

The scolcx is ciubsnaped^according to Barrois and 
Marotel, spheroidal according to Schwarz. The figure or tne head 
as aeiin3ated oy Marotel and Sohwarz are reproduced (Pi. rigs )« 
Barrois and Scnwarz reported that there was no rirth sucker and 
no rostelium while Marotel tnougnt that the slight elevation at the 
middle or tne head.was a rudimentary rostellun. He saw no fifth 
eucker. Marotel states that the globular suckers are directed 
anteriad. The head (PI. 7 rig. Ijf ) which I observed was cleared 
in glycerine. It presen sa somewhat tetragonal race., with a small 
elevation at its center which is not a rostelium and upon which 
there is no flth sucker. The -suckers are placed in the corners 



oi the anterior lace and they are directed anteriad. Slight grooves 
or wrinkles wnich do not extend to tne apex partially divide the 
head into quadrants. Tne suckers are nearly round in outline. They 
entirely lack any trace oi The inturned lower margin which is 
found in The suckers oi Crepidobothrium Gerrardii * They measure 
u.£y-u.4_b mm. in diameter with a sucker opening u.i 7-0 .lbb mm. in 
diameter. The sucker cavity is deep. The head is 1,19 mm. broad^, 
O.bO mm. long^O.y^b mm. thick. The following comparative table 
presents The data on The nead, secured by Barrois, Llarotel, 
Schwarz, and myself. 









Olub shaped 


somewhaT N 

breadth i-l<#3 
" 1.U5" 

" 1.0 " 

i.iy " 
length O.bO " 


round in 

u.3£-u # 34 mm -0.34mm 
0.3 l J-:. .408" 

u ,120mm 




Barrois (idyb: 

MaroTei (lbyy; 

Schwarz (iyo8: 
24 j 

La Hue 


mm. i 

I found That The neck was about b-b mm. long by u.ybu mm. 

broad. Its limits could not be readily determined in a Glycerine 


preparation. Uarotel state s^ the neck is 4 cr b mm. long and u.bbu 
mm. broad at its narrowest place. Barrois and Schwarz do not state 
its dimensions. 


The first proglottids are much Droader than long. These 
Increase in length more rapidly than in breadth. Marotel ( luyy ) 
states that at 2u mm. rrom the head the proglottids measure 0.255 
mm. long by u.b3 mm. oroaci^at nb mm, o.b5oxo.b5. mm. i. e., 
qutdratej at 30 mm^ 0.710 mm. long^0.67U mm. broadj at 40 mm., 1.45 
x 0.75 mm. and the iast proglottids are 3 to 4 mm long by 1.0 mm. 
broad. In my material nearly mature proglottids measure as muoh as 
o.y mm. broad by 0.55 mm. long and ripe proglottids about 0.85 mm. 
square or at times a little longer than broad. Ripe proglottids 
measure z-6 mm. long by 1.0-1.2 mm. in breadth. In s^me contracted 

: regions ol' the strobila the breadth may measure as much as 2.0 mm. 

GENITAL 0RGA1T3.- These are typical or the genus. The 

; sexual aperture is irregularly alternating, situated, in mature 
proglottids at, or slightly; anterior to the middle of the margin, 

i v;hilo in ripe proglottids it may be either anterior or posterior 
to the middle. Both cirrus and vagina open into a common genital 
sinus v/hioh is very snaiiow. The vagina lies anterior or posterior 
to the cirrus-pouch with almost equal frequency. In ripe proglottids 

\ frequently and in mature proglottids more rarely, the vagina may 
open dorsal to tne cirrus-pouch. ^ p/.tf/-"^^*/ 

I-Iale organs . - The testes (PI. 7 i'igs.fP^J^ number from 130 
to Ibu in my preparation and those also are about the limits 
pbserved by the other investigators. They measure from 0.026-0*037 
mm x 0. 053-0 .Ub3 mm. in my preparations. Others report them as 


measuring U.Ob x u.U4 ram., u.Ub^-U.UYii mm, and x U.U4 mm. Tney 
| are arranged, in two tz elds wnicn in elongated proglottids (PI. /Tig. 
^are near the ventral excretory vessels while in contracted prog- 
lottid si PI »7 tl&fti ) they are some distance from the ventral 
vessels, in the former case the testes or each field are arranged 
in two irregular rov/s wnile in the latter case the testes ol' each 
field are irregularly arranged in a broad fjeid. The testes are 
dorsal to the uterus. 

The vas deferens (Pi. 7 llgJ3? ) forms a mass of coiis 
which extend from the cirrus-sheathe nearly to the middle of the 
proglottid . My drawings (rl.7 i'igs./32i/#] show it more heavily 
developed than do the figures of Mart ot el (itiyyj which are repro- 
duced ( tigs M IC *} • 

The cirrus-pouch is relatively short and broad, 0.26- 

0.2'JO mm. long by 0.102-0.136 mm. broad. In the broad mature and 

ripe proglottids it extends but a short distance within the vitellariii* 

The ratio of its length to the breadth of mature proglottids is 

| about 1:6 , to the breadth of ripe proglottids about 1:5-1:55, to 
i ■ * 

j the breadth of the ripest proglottid observed about 1:4. Schwarz 
states that the cirrus-pouch is about 0.33 mm. long. 

The ductus e jaculatorius (PI. 7 fig./£* } has numerous 
coils. The unprotruded cirrus is broad and muscular. I have not 
seen it protruded and none of the other investigators report having 
seen it protruded. 

An examination of Marotel's figures which have been 


reproduced (Pl.^f ±lgsM® shows fewer coils oi" ductus e jacuiatorius 
and -oi' vas deferens than do mino (PI. 7 figs./H/#). This can only 

' be explained by the assumption that he could not follow these out 
completely in his preparation which he states were made from material 
in a very poor state of preservation. The material which I worked 
upon was in a fair state of preservation and these ducts could be 
traced with ease. 

Female organs .- The vagina (PI. 7 f : gsJ#$ at its distal 
end is very hroad , ram., which is due in part to its 
broad lumen and in part to its thick sphincter vaginae. In this 

i region the vagina is nearly as broad as the basal portion of cirrus. 
The vagina in its course to the middle of the proglottid never crosses 
the cirrus-pouch. Arrived at the middle^ it bends sharply and takes 

' a sinuous course to tne interovarial space which it enters from 

j the dorsal side of the owary not from the ventral as Marotel figures 
it. There are no coils in the vi gina anterior to the ovary. A 

i receptaculum seminis has not been observed. 

In nearly mature proglottids the lobes or the ovary are 
more slender and more pointed at the tips than in mature and ripe 

! proglottids. In well elongated proglottids the lobes of the ovary 
assumes the shape figured by Marotel (Pi. ft" f lga«/l£ft) • 

The vitellaria are follicular. The follicles are small. 

i The viteii iri9 ducts cross the o V arys ventrally instead of lying 


some distance anterior to them as Marotel figured them. The uterus 
is a median "cube in mature proglottids. The early development of the 
lateral pouches (PI. 7 may be traced as small, somewhat 

excentric bulbous^ enlargements oi" the median tu<>e. This is somewhat 
differant than in 0. f ilaroides and 0. perspicua . These enlarge- 
ments may attain same size before any eggs are developed. In the 
early stages there is considerable resemblance to the condition 
shown in Marotel's figure (reproduced in Pl.^ fig./^yfe} of the 
vigerous segment of 0. Calmettel . Later however many of these 
pouches extend toward the sides and the uterus resembles that of 
other species of the genus. The wails of these pouches are not 
very easily observed so Marotel may have overlooked them in his 
specimens. The pouches' number 24-35 on either side. Two to four 
or five ventral uterine pores were observed in some of the riper 

EGGS.- The uterine eggs in the alcoholic material are 
surrounded by three membranes. The outermost very thin, hyaline 
meraDrane measures from u.024-u.u2t> mm. The second, thick a.nd 
homogeneous, membrane measures U.022 mm. The inner membrane is 
thick, granular and more or less irregular in outline. It measures 
U.U14 mm. while the six-i.ooked embryo measures u.012 ram. in diameter, 
Marotel (lyyy) states that the eggs are globular, uith two raernbranes# 
The outermost one is -cnin and membranous, u.Ubb mm. in diameter. 
The otheryiomogeneous and somewhat thick^is U.U24 mm. in diameter. 


This merabrano corresponds in size and description to the one wnich 
I call the middle membrane • He further states that the embryo 
is granular but does not give its dimensions. 

RELATIONSHIPS.- This species varies from the 0. racemosa 
described by Schwarz (iy(J8) in the much larger size of the head, 
the larger size tit tne suckers, and the relative prominence of the 
same. The number and arrangement of the testes are radically diff- 
erent. In the length of the cirrus pouch, in the number of coils 
of ductus e jacuiatorius they differ greatly. In the size of the 
vagina tnere is considerable difference • The character of the 
diverticula of the uterus is not the same. In this species the 
uterus extends back to tne ovary while in 0. racemosa as described 
by Schwarz it does not. The vitellaria are much alike in 

This species is much smaller than 0. gr audi s . The heads 

are of about tne same size but not alike in shape. The relations 

and vagina 

oi" cirrus, eirrus-poucn^ are different. There is a wide difference 
in the number of testes and in tne number of uterine outpocketings. 
The hosts and geo/rrapnical distribution are also widely different. 

It varies from 0. Mai enzelier i in having a much smaller 
head, smaller suckers and in tne lesser prominence of tho same 
its smaller proglottids and in itz smaller number of testes. 
The relations of the cirrus and cirrus-sneat .e are quite different 
The fact that the vagina lies usually posterior to the cirrus- 
pouch in 0. Marenzelleri constitutes a marked difference. 


The shape oi' the ovaries is somewhat different. In^, Marenzeller i 

^marked "bending of tne vagina in its course to the middle of the 

proglottids is very unlike the condition in thr's species. 

0. Calmette i is larger than 0. t rimeresur i. It has a 


larger head, larger suckers and^ than twice as many testes. The 
relations of the unprotruded cirrus and the cirrus-pouch are much 
alike in the two species. 

0. Calmettei is so much larger than 0, perspicua^ O. 
Natter eri or 0. pigmentata • That any possiolity of confusion with 
them is precluded. It is 1 ike wise much larger than 0. f ilaroides 
and 0. Lonnborgii which occur in Amphibia. 

0. Calmettei most nearly resembles 0. g rand is 
0. Marenzelleri and 0. trimeresu ri which likewise occur m the 



[P/Jrf>'?3.1??- 1 37 TP/- V. f<P«f-/5y 
ltfU8: Taenia trimoresun Parona I8y8: 7-il. 

I8y8: Icnthyotaenia t rimeresun Luh© l8y8: 6b2. 

1 V J08: Icnthyotaenia trimerasun Schwarz iyoti:33-3b. 

SPECIFIC DIAGNOSIS.- Characters of genus. Length up to 
lob mm. Llax breadth o,7b-l.b mm, Scolex unarmed, without rostellum, 
broader than neck, diameter 0.7b mm. Suckers prominent, hem- 
ispherical, muscular, O,026-u,2b mm. in diameter, situated anteriorly* 
No fifth sucker. Neck three times length or head, t;2b-u.b mm, broad, 
First proglottids broader than long. Mature proglottids quadrate or 
longer tnan broad. Angles of proglottids nox pro uinent . Strobilation 
indistir. ct . 

GENITAL ORGANS.- Genital pore situated near middle of 
proglottid margin, irregularly alternating. No genital papilla. 
Genital sinus if present very shallow. 

Hale o rgans.- Teste5 1UO-108 in number^ measure ing j.0b3 
x U.U27 to U.U8U x U.U27 min, r two fields not near .iteliaria 
T stes not posterior to ovary. Mass or coils of vas deferens not 
larger. Ductus e jacuaitcrius much coiled. Cirrus muscular. In base 
of protruded cirrus many coils of vas deferens. Cirrus-pouch about 
0.27U-U.330 mm, long x 0,i3b mm. broad. Cirrus-pouch into proglottid 
breadth 2 1/2-4 -cimes. 

Female organs , - "^agina anterior or posterior to cirrus- 
pouch ,not crossing same. Lumen oi vi.gma varialbe in size. 


vagina present. Lobes or ovary flattened^ elongated, narrow. 
Vitelline follicles small. Uterus when fully developed with 2u- 
3U (?) lateral pouches. 

KGGS.- ITo ripe eggs observed. 

HABITAT.- in the intestine of T rine re sum s f orm ftsus 
( type hostj ^Island or MentaJiei , East Indies (type locality), 

TYPE. Specimens in Parona 1 collection labelled 
" Aryncno taenia trimorasuri Par., 'i' rimeresur us rorm$losus . ( Montatt fei ) 

Also two toto preparations oi same material in collection Ifrof, 
H. B. Ward. 

HISTORICAL SUMMARY,- Parona (18y«:7-llJ described and 
figured this species wnich occurs in a Trigonoc eprt id , Trimoresurus 
formos us . 

Luhe [ ltiy&:bi>2) stated that this species belonged, to the 
Ichthyotaenia . In a footnote he says Herr Prof, Parona hat mir 
die Originalrexemplare der art in unei.^ennutzigster Weise zur 
beifugung gestellt, so dass ich die Zugohoiigkeit derselben zu 
der fiattung Ichthyotaenia feststellen konnte." 

Schwarz (1^08) who had no specimens for examination re- 
wrote Parona's description but was able to add nothing new . He 
believed this form to be a spceies of Proteoce phalus and called 
attention to the part that the lower part of the uterus in Paroni's 
drawing was probable the ovary and that the testes would be found 
in the anterior regions of the proglottid where they were frequently 
very difficult to see. 

MATERIAL. - Thanks to Dr., H. B. Ward I have fiad some 


o±' this i'orm from Parona' s collection . The material consists 

o±" about a half dozen pieces among which no head could be found. 

This material is labelled A rynoho taenia trimeresuri Par, 

Trineresuris f ormosus ( Ment awel ) . Prom it two toto preparations were 

made. These together with the alcohol jcs and Parona 's original 

work from the basis 01 the following description. 

EXTERNAL CHARACTERS. - In the material at my disposal 

I found no head, neck or first proglottid^. The peice^ measured 

10-20-69-7$ mm. in length by a max. breadth of about 1.0 mm. Parona 

in a table gave data concerning the length and breadth of five 

specimens with heads and five without. The length of those with 

heads ranged, from 12 to 50 mm. by a max. breadth of 0.75 to l.o mm. 

while the specimens ./ithout he £dS measured from 33 to 105 mm. in 

length- by a raaz.> breadth of .76-1.5 mm. 

The scolfrx (Pitf fi£^ reproduced from Parona (i8yu)^ 

is spheroidal, is unarmed and without 

roitcv'lum and oroader than the neck. Its diameter is about 0.7b 

mm. The suckers are prominent and situated in the anterior part of 

the head. They are hemispherical, strongly muscular and are 0.16- 

U.25 mm. in diameter with an aperture of u. 11-0.16 mm. 

he nec:*: is aoou-c i,hree times as long as the scolex 

Its breadth varies irom 0.^6-0.5 mm. Thf* first proglottids aro broader 


than long but not very short. The^ following gradually became longer, 
the last being longest. However, the increase in length is not 
constant for short proglottids are found between long- ones, n< r 


is the "breadth ol' the strobila constant. Some proglottidsare 
about quadrate, otners are 2-3-4 times as long as broad. 
The proglottids limits are not pronounced and the strobilation 
is not distinct. The angles of the proglottids are not prominent. 
Hence the worm has the appearance of a continuous ribbon. The 
i'oregoing description or the head, sucker^neck and proglottids is 
based on the original work of Parana (l8 l J8:8-y). 

In the toto preparations which I examined were a few 
mature proglottids. One proglottid (Pi. J figJJJ) contained a 
few eggs in tne uterus. These were omitted from the drawing. The 
length of tnis segment was 1.7 mm. and its breadth O.y mm. It was 
thin and flat, tho considerably thicker than proglottids of a 
• similar degree of development from 0. perspicua or ffattereri 

GENITAL OR GAIT 3. - The genital pore is situated near 
; tne middle of the proglottid margin. It alternates irregularly in 
the strobila. The pore is not marked by a papilla, nor by a deeply 
indented region as in 0. grandis . The vagina and cirrus-poucn open 
very near each other. The presence of a genital sinus is difficult 
to determine from my toto preparations. The drawings fPl. figs ) 
do not snow one. If present at all it is very shallow. 

Mais organs.- The testes (PI. J fig./? 7) number 
about 100 to 108. They measure from 0.UG3 x 0.027 mm. up to 
o.OtfO x 0.u^7 mm, the long axis of the testis being perpendicular 
to the long axic of the worm. The testes are arranged in Lwo 
narrow bands. Well away from the vitellaria. In this respect the 


ohe form resembles 0. Calmet tel 

Hone of "Che testes are posterior to the ovary tho Parona's drawing 

which has oeen reproduced (PI. IT fig./Yf ) snow them there. 


The vas deferens (PI. 7 fig«/3 7 ) a small mass of coils 
in -che mid-region of xne proglottid. The vas deferens after 
entering the cirrus-pouch becomes the much coiled ductus ejacuiatoriuii 

( PI . 7 Z±g$,H5>l3(o) . 

The ductus e jaculatorius passes over into the thicker 
walled and more muscular cirrus. Under certain conditions the 
cirrus is protruded (PI.?" f igs./3-£/37) . In this case the basal part of 
cirrus is dilated and the greater part of the ductus e jaculatorius 

is crowded out into the dilated "basal part ( PI./' f ig. Of) . 

A more highly magnified drawing of this cirrus and vagina is 
shown l^i PI, 7 flg./3^Z This condition of the cirrus is very 
similar to that described and figured by Schwarz (iyu8) for (V 

! Marenzelleri . Schwarz, however, claimed that the whole cirrus-pouch 
was evaginated. In that he misinterpretted the facts for only the 

, cirrusand a part of the ductus e jaeuiatorius are pushed out. More- 
over his own drawings which are reproduced (PI. % fig./^y) snow 
tne cirrus-pouch in its proper position. In Schwarz's drawings of 

I 0. Marenzelleri the cirrus beyond the base is filiform. That 
condition has not oeen observed by me in this species but it is 
highly probable that at least a part oi the coils within the broad 
basal portion of the cirrus may at times be pushed out to form a 
filiform cirrus. Unless this be the case it is difficult to under- 
stand how copulation can be possible. 


The large number o±" coils of ductus e jaculatorius in this 
species furnishes a differentiating cnaracter between this species 
and 0. Marenzeller i. 

The cirrus-poucn is large and very muscular. Its length 


varies from 0.27U-U.34U mm. It is about 0.13b nmi. breadth. Its 

length according to Parona is 0.33U mm. The ratio of The length 

of "Che cirrus-pouch to the breadth of proglottid is from 1:4 to 2:5. 

The vagina lies either anterior or posterior to the cirrus- 

Female organs .- Near its opening. (P1.^7f igs,tf%) it 

has a very heavy sphincter muscle. The vagina does not cross the 
. irrus-pouch. Its course after reacning the median field is fre- 
quently quite sinuous. In soae parts or its lumen tne vagina is 
narrow wnile in otner regions it may be widely dilated. 

The lobes of tne ovary are flattened. They are more 

elongate and narrow A than in 0. Marenzelieri . The prired vitelline 
ducts cross the ovary on the ventral side. 

Tne uterus in mature progiottids is a median tube extending 
tne lengtn of tne progiottids. No ripe progiottids could be examined 
so the number of lateral uterine . outpocketiugscouid not be determined' 
However, tne drawing (PI. 7 f"ig./?7 ) indicates that probably about 
2u-ao lateral poucnes would be developed, on eitner side. 

EGGS.- No ripe eggs were present in tne material "examined. 



The description drawing 01 xne manure proglottid 

shows this to be a Proteocepnaiid species which rails readily into 

the group infesting snakes. My description and drawings no not 

agree very well with Paronas except as to size, position of genital 

pore and length and shape of cirrus-pouch. From his drawings it 

appears that he has confused certain organs. The lower portion of 


the uterus in his drawing^undouotedly the bilobed ovary. The 
position of the testes posterior to the ovary in his figure cannot 
be explained except on the supposition that they are parts or the 
organs or the inter-ovarial space. The sinuous duct which he shows 
as the vas deferens extending Dacic from the cirrus-pouch to the 
ovary is not the vas deferens Dut the vagina, which in fact passes 
below the coils of vas deferens before reaching the middle or the 
proglottid. The mass or coils or the vas dererens is poorly shown 
at the inner end or the cirrus-pouch. The opening or the vagina, 
I the vaginal sphincter, the first part of vagina thru the length of 
the cirrus-pouch he has omitted altogether^ *he has confused it with 
a part of the cirrus-pouch. The cirrus-pouch as ho drew it seems 
' to be made up of lamellae. In a poor preparation the cirrus-pouch 
right have had this appearance. The cirrus within the cirrus-pouch 
is not properly delineated. The uterus in an early stage may 
possibly have had the single anterior lateral pouch on either side 
as he has shown it. I have not seen a proglottid as nearly ripe 
as tiiis one is. The testes and vitellaria he has omitted from 
his drawings. It seems highly probable to me that Parona's drawings 


are intended to represent tne same species as my drawings do. The 


apparent difference came about tiara Paroras using poor pre par ax ions 
i tor study. 

RELATIONSHIPS.- 0. trimeresuri Parona in size is mucn 
smaller tnan 0. grand! s . 0. Harenzelleri and C repidobothrium 
Gerrardii . Moreover in number and arrangement or testes > size and 
proportions of cirrus-pouch and cirrus tnese tnree species differ 
from 0. trimeresuri . In size tftis species is more nearly related to 
0. Calmettei but in the latter species the relations or tne cirrus, 
cirrus-pouch and vagina are different. Tne testes in 0. t r imeresuri 
are arranged mucn as tney are in 0. Calmettei but the size of the 
head and the suckers are greatly different in the two species. _0. 
Nattereri and 0. perspicua are smaller, more delicate and have very 
different relations or cirrus, cirrus-poucn, vagina, and testes. 

This rorm dirrers rrom any other species tftus far &eseri£- 
afrrom snakes yet it has its closest affinities with 0. Calmettei . 
liiiew se rrom a Trigonocepnalid. 


(PI. % figsJ7W) 

1019 : 

Taenia racemosa 

Rudolphi , 


: u9j. 


Taenia ooluto*i 

Rudolphi , 


: 709. 


Taenia racemosa 



: 610. 


Taenia racemosa 



: 511. 




Ichthyotaenia racemosa Barrois , 

1898: • 



Ichthyotaenia racemosa Liihc, 





I eh thyotaenia racemosa Schwarz, 




SPECIFIC DIAG1I0SI3.- Characters of genus. Length of 
strobila up to 150 mm., breadth about 1mm. length of ripe proglottid 
1 ram. breadth 1 mm. 

Scolex easily distinguished from strobila, breadth of 
scolez about 0.5 mm. Suckers four, nearly circular, Q.f:7-0.30 mm. 


in diameter. 

GE1TITAL OR GAITS :- As in genus. Testr not in two distinct 
fields. Tests 0.072-0.078 mm. in diameter, 100-120 in munber. 
Ductus ejaculatorius sinuous. Cirrus-pouch reaching nearly to 
middle of proglottid. 

Vagina anterior or posterior to cirrus-pouch. Ovary 
voluminous, lobes almost v.inglike. Uterus with about twenty long 
out pocket ings on either side. 

EGGS.- Eg^s with three shells. Diameter ci egg 0.024 mm. 


HABITAT . - In intestine of South American Snakes. 





Coluber sp 

Ophiomorphus miliarls 
Ophiomorphus miliaris 

Ophis Merremii 
Ophi 3 Merremii 


Museum Vienna 
it it 


batterer Rudolphi 1819:692. 

Diesing 1850: 511 . 
Schwarz 1908:28. 
Diesing 1850: 511 . 
Schwarz 1908:28. 


HISTORICAL SUMMARY.- This species was first described 
>y Rudolphi (1819:692). Since this description is not readily 
accessible to many workers I quote in full: 

Taenia racemosa R. n. sp. Pone Synops. n. 52. 
T. Gapite oboonico, collo brevi, angustissimo , articulis 
planis, elongatis, forminibus marginalibus alternis prominulis. 
Hab. In intestinis Colubri n. 20. specimens sexpollicare et fragment a 
hujus Taeniae HAfflBEEBR in Brasilis reperit. 

Caput antrorsum dilatatum, sive obconicui, osculis 
orbicularibus , aut hemisphaericis , nam satis profunda videntur, antiei||s 
Collum breve, angustissimum. Arti ouli . quos vidi, plani, tenues, 

elongati, lineam ad sesquilineam longi, tertia lineae parte latiores, 
foraminibus marginalibus alternis, prominentibus. 

Ovaria singulorum articulorum lineam mediam fere totam 
sibi vindicant, ad cujus latera utrinque maculae exiguae opacae ita d 
digestae sunt, ut ovaria racemosa appareant. 

Obio . Caput Taeniae ormphalodis Synops. n. 9. articuli vero 
T. tuberculatae n. 25. ut fragments, quae vidi, Taeniam illis interme&V 
iam reddant. 

Rudolohi (1519:709) established the name Taenj a Colubri 
to designate a few proglottids found in Colubri . Sp. , Brazil by 
llatterer, I quote: 

Taenia Colubri. Pone Synops. n. 140. 

Fragraenta in intectinis Colubri n. 12, a HaTT_;REP0 in 
Brasilia reperta Museo Viennensi debeo lineam circiter longa, 
duodecim ad octodecim articu is constantia, latoiribua quam longis, 



subcuneatis, angulis plus minus exstantibus, capite destituta. 

Ut'rum vere colubrina, an ex ave quadam deglutita forsan 


Dujardin (1845:610) added nothing to Rudolphi's description, 
Diesing (1850:511) gave a short diagnosis of a form from the museum 
at Vienna which he identified as Taenia racemosa Rud. His descrip- 
tion varies so widely from that of Rudolphi's and his list of hosts 
covers such a range of families that I quote them in full: 

^ Taenia racemosa RUD01PHI. Caput magnum tetragonum, 
aoetabulis angularibus subterminalibus V. terminalibus , subovatis V. 
cordatis. Collum nullum. Articuli supremi brevissimi, subsequentes 
longiores et latiores, angulis retundatis, ultimi longi parallel- 
opipedi angustiores. Penes filiformes basi incrassati, marginales 
vage alterni. Longit. 2 — 1-J-; latit. med. 1 — 3 ; ultim. 1 

Taenia racemosa Rudolphi : Synops. App. 692 — Dujardin : 
Hist. nat. des Helminth. 610. 

Habitaculum. Ophiomorphus miliar is . Martio. — Eunectes 
Scytale . Octobri, Februorio et Junio. — Bothrops Mararacca, Mertin 
et Aprili. — Ophis Merremii. Januario, in Brasilia ( flatterer ) : 
in intestinis. M. C. V. ' 

Von Linstow (1878:183) did no more than refer to Rudolphi 
(1819) and Diesing (1850). 

Barrois (1898:3) was convinced that Taenia racemosa Rud. 

belonged to the genus I chthyo taenia for he had examined a head and 

some ripe proglottids "provenant du type primitif de Rudolphi." He 

did not describe the species. 

Luhe (1898:652) stated that Taenia racemosa belonged to the 
L I 


genus Ichthyotaenia. Marotel (1899:34) quoted Die sing's (1850) 
description or this species which he stated was too incomplete to 
permit a precise determination. 

Parona (1901:3) reported some specimens collected "by Dr. 
Adolf Lutz from Bothrops ( Lachnis ) lance olata , S. Paulo, Brazil under 
the name Ichthyotaenia racemosa Rud. These specimens which have been 
examined "by ne prove to "belong to the species Ophiotaenia Calmettei 

Shipley (1905:101) reported some specimens of Taeni a 
racemosa Rud. from Euqectes murinus Wagler. Prof. H. B. Ward se- 
cured some specimens "bearing this name from Mr. Shipley. Upon 
examination they proved to be specimens of Crepidobothr i um Serrardii 
(Baird) . 

Qchwarz (1908:28-£9) secured some material which Barrois 
had obtained from Dr. E. von Marengeller of Vienna. He also secured 
additional material from Dr. E. von Marenaeller. Upon this material 
he beased his description of Taenia racemosa Rud. He did not de- 
finitely state the name of the host from which his material was 

DISCUSSION.- It is possible that T. colubri Rud. collected 
by Batterer in Brazil from Coluber sp is a synonym of T. racemosa 
Rud. also collected by Batterer in Brazil from a special of Coluber . 
Rudolphi's description of T. colubri i s insufficient to place the 
species. Diesing (1850:558) states that T. colubri was collected 
by Batterer in Brazil from Ophiomorphus poecilogyrus . He did not 
give any descriptive data. 


Diesing's diagnosis of Taenia racemosa Rud. does not agree 
with Rudolphi's discription of that species. Diesing's diagnosis 
in regard to the tetragonal head, the terminal or subterminal 
angular suckers which are subovate or cordate fits the diagnosis of 
Crepidobothrium Gerrardii but not of P. raoemosus. The latter 
species has orbicular suckers. Likewise Diesing's statement that 
there is no neck, that the first proglottid are very short, follow- 
ing ones larger and broader, with rounded angles the last proglottid*; 
long parallel opipeds and somewhat narrow agree much better with the 
descriptions of £. Gerrardii than with that of P. racemosus . 
Diesing's form was considerably larger than Rudolphi's. Ho complete- 
ly protruded cirrus was noted in my material of C_. gerrardii hence 
the cirrus of the two forms cannot be compared. 

Moreover Diesing lists Eunectes scytale as a host of 
Taenia racemosa . Eunectes scytale is a synonym of Eunectes murinus . 
a species in which £. Gerrardii but no other Proteocephalid has been 
found. It seens therefore that Diesing's diagnosis of Taenia 
racemosa was based on the material from Eunectes infer inus : ( scytale ) » 

A further analysis of Diesing's list of hosts of Taenia 
Racemosa shows that besides the Boidae , represented by Eunectes 
murinus . the Colubridae are represented by two species and the 
Crotalidae by one species. It seems probable that Diesing's Taenia 
racemosa must have included several species, for among the species 
of Ophiotaenia from snakes which have been adequately described in 
recent years no species has been found in hosts belonging to differ- 
ent families. A list of these species, their hosts, and their 


distribution is found elsewhere (P.). 

Another fact to be remarked about these Proteocephalid 
species from snakes is the size of the individuals of that species 
which infests the Boidae is larger than that of the species which 
infost the Grotalidae and these in turn are larger than tho::e that 
infest the Colubridae . By the term size T mean ~ize of head, 
size of suckers, breadth of neck^breadth and length of proglottid^ 
and length of strobila. In certain of these characters my general 

I statement i^ils but, considered in a broad way, it is tnue. This 
is additional evidence that Diesing included several species under 
the one narce Taenia racemosa . 

Concerning hosts of Taenia racemosa Schwarz (1908:28) 

^ says: " Als wirte werden angegeben: O phiomoiohus miliaris . Ophis 
Merremii . Bothrop s jararacca und Sunectes scytale . Schlangen, die 
in Brazilien vor'-oramen". This seems to be but a restatement of 
Diesing' s list of hosts. Nowhere does Schwarz state definitely the 

' exact host from which his specimens were taken. 

A study of his description in comparison with other species 
1'rora snakes shows that his specimens are most closely related to the 

I species from Colubridae yet they are distinct from any of those 

' species. It seems then quite probable that his specimens cane from 

j one of the Colubrid, e . possibly OphiomorphuL nr'l iaris or Ophis 

The emended I ist ol hosts of Taenia r acemosa thon contains 
these names, Coluber sp., Qphioraorphus miliaris and Ophis Merremii. 


Further questions regarding Schwarz's T aenia racemosa 

arise. Is his species identical with Rudolphi's Taenia racemosa ? 

Could Schwarz have had any of Rudolphi's type material? At my 

re:ue; t Professor ".Yard wrote to Proiessor E. Marenzeller at Vienna 

for information concei-ning the specimens of Taenia tl cemosa which 


he had sent tc Barrois to Schwarz. Professor Marenzeller replied 

that all of Rudolphi's types were in the Museum at Berlin. It is 

theref one impossible that Schwarz or Barrois had Rudolphi's type 

specimens for examination and the question of the identity of 

Sch arz's Taenia racemosa and Rudolphi's species of that name 


remains open and must so remain until Rudolphi's t; pes ^re-examined, 
if they still exist. Meanwhile I shall assume that the Taenia 
racemosa of Schwarz and Rudolph; are identical. 

I.IATERIAL.- Schwarz's material all come originally from 
the Museum at Vienna some directly through von Marenzeller, and 
some through von Marenzeller to Barrois, then from Barrois to 
Schwarz. As hosts he mentioned four species of South American 
snakes two of which have been ruled out in the discussion leaving 
Ophis Merremii and O phioracfcohus mi liar is as probable hosts. If 
the specimens v/ere from Ophis Merremii they were probably collected 
by batterer in Brazil. 

The following description is based on the W' rk of Schwarz 


EXTERNAL CHAI-.ACTURS . - The observed length of specimens wa; 
160 mm. he length oi ripe proglottids 2 mm. and the breadth of the 
same i mm* 


The scolex^is easily distinguished from the strobila. 
It bears four nearly circular suckers which measure 0.:l70-0 .300 mm. 
in diameter. The scolex is 0,, 40 mm. in diameter. 

"EHITAL ORGANS.- These^ agree in arrangement with the plan 

of other Proteocephalid;- infesting snakes. 


The testes are A exceptional size and appear as large spheres. 

They are not limited to the side fields "but are scattered irregularly 

through the whole segment. The testicular field is not interrupt d 

at the anterior or posterior margins of the segment. The testes 

number about 100-120 and measure 0.072-0 ,07 J mm. in diameter. 


The ductus e jaculatorius has a sinuous course^ the cirrus- 
pouch but makes no coils. The cirrus-pouch lies perpendicular to 
the longitudinal axis and reaches almost to the median line 

Female organs . - The vagina opens anterior or posterior 
to the cirrus . The ovary is voluminous. It is bilobed, 
joined at the mid le of a mid-piece. The two lobes are of a 
somewhat plumper form than are those of B. Marenzelleri . The uterus 
forms on ither side t about twenty long slender diverticula. 

EGGS.- The eggs are round and have three menbranes. 
Hooklets are not presnnt on the outer meHbranee. Diameter 0.024 ram. 
Schwarz does not state whether this is the diameter of the whole 
egg or of the emhryo. 


RELATIONSHIPS.- Proteocephalus racemosa is a much 
smaller species than Crepidobothrium Gerrardii. The head and 
suckers are smaller, the proglottids shorter and narrower. The 
form of the suckers is greatly different in the two species. 

p_. racemosa differs from Salmettei in its smaller head, 
smaller suckers, in the distribution of testes and in the number 
of uterine pouches* 

0. racemosa is likewise much smeller than 0_. Marenzelleri 
and 0. grandis n. sp. in length, breadth, siae of head, and size of 
suckers, it also has a fewer testes . 

£• racemosa differs from 0. trimeresuri in being somewhat 
smaller. Its chief differences lie in the position of the testes 
and in the character of the cirrus and vagina. 

racempsa differs from 0. pigmentatus in having larger 
suckers. 0. pigmentatus is too poorly described to furnish more 
diagnostic, differences. 

0. racemosa differs from 0. )lattereri and 0. perspicua in 
having a larger head, larger suckers, a different arrangement of 
testes, and smaller proglottids. 


OPHIOTAEHIA PtOflCA. ( Cholodkovsky} 
(PI. f FlgB../Sftf# 
iyu8: Taenia punic a Cholodkovsky 1908: 419-20 
1'JIO: Proteooephalus punicus Hall 1910: 148. 

SPECIFIC DIAGNOSIS.- Characters ol' genus. Length up 
to 10 cms. Breadth up to 2.7& mm. head large, 1.5 mm. b road, 
V.v.A.t'wied, somewhat tetragonal. Suckers rour, large round, 0.7 mm. 
i.. ^a-.. diameter. Sucker musculature heavier on inner half. Neck 
short. Segmentation not evident. Proglottids attached by full width. 
First proglottids "broader than long, older ones longer, quadrate, 
and finally longer than broad. 

&EIJITAL 0RGA1JS.- The genital opening alternates 
irregularly, situated near middle of proglottid margin. 

Hale Organ s.- Testes very numerous about 200, situated 
in two well defined lateral f^lds. Cirrus-pouch siedner, extend- 
ing from 1/4 to 1/3 across the breadth of the proglottid. 

Female organSj.- lobes of ovary long, slender, rough in 
outline, connected by long slender midpiece. viteilaria lateral 
composed oljsmall follicles. Uterus median. 

Eggs.- Hot observed. 

Chlolodkovsklf ( ±yoy :4ia-2o ) reported this soeaies from 
a dog in Tunis ( Icland ol DschorbuJ. Since this is peculiar host 
in whsa\h to find a species normally occuring in a lower voretbratc 


I give lis description omitting only the first and last paragraph^ 



Nieht ohne TTberrasohiing habe ich also in einer mir von 
Dr. LI. Vieiyi&cTQ zugesandten Tanie aus dcm Darme eines Hundes Eine 
neue Species erkannt , die ich hier in aller Kurze beschreiben will, 
Das Material (in Formol aufbewahrt) stammte aus Tunis ( inscl Dsch- 
erba) und bestmd aus drie unvollstandigen Stobilae, deren 
groBte eine Lange von etvra 10 cm erreichte. Der Scolex ist sehr 
groB (1,5 mm breit), unbewaffnet, mit vier starken rundlichen 
Saugnapfen and einer kleinen Erhobung auf dern Scheitel versehen. 
Die Saugnapfe sind etv;as asymmetrisch gebaut , indem ihre innere 
Halfte meklich dicker als die auBere ist; der groBte (auBere) 
Durchmesser derselben betragt 0,7 mm. Gleich hinter enim sehr 
kurzen Halse wird die Strofeila etwa 2 mm, dann bis 2,75 mm breit, 
nach hinter aber allmahlich etwas enger. Eine auBere Gliederung 
ist auf ganzen Stucken unbemerkbar und laBt sich erts auf gefarbtei. 
Balsam- oder Glyzerinpraparaten in der Gcstalt von sehr schwach 
ausgesprochenen Querrinnon konstatieren. Die ersten Proglottiden 
sind breiter als lang, dann v/erden sie allmahlichlanger , quad- 
ratisch und endlich langer als breit; in keinem der mir vorgelegen 
en Exemplare waren aber ganz reife, d.h. fertige Eier enthaltende 
Proglottiden vorhanden. Die Geschlechtsof fnungen alternieren 
unregelmaBig. Die stark in der Querrichtung verlangerten Oyarien 
ligen dicht an der hinteren Grenze der Proglottis, der Dotterstoel: 
ist sehrklein, der Uterus bildet einen geraden, in der Llittellinie 
der Proglottis nach vorn verlaufenden Stamm. In vollig reifen 
Proglottiden triebt er vielleicht auch laterale Zweige, in meinen 
Examplaren v/ar en aber immer einfach. Die zahlreichen Hoden ligen 
in zwei lateralen Lang£;f eldern nach inr.en von den groBen Encretiom 
gefaBen. An der Stelle der logzteren befindet sich in totalen, 


mit Boraxkarmin tingeirten Praparaten der reifsten mir vorgelegen 
Proglottiden eine kornige Liasse, deren Bedeutung mir unklar geb- 
lieben ist, da ich aus Mangel an Material keine darauf bezuglichen 
Schnittserien verftigen konnte. 

Da die hier beschriebene Tanie alien Anschien nach eine 
neue Species darstellt, so schlage ich vor, sie nach ihrem Fundort 
•Taenia punica zu nennen." 

This species was not noted by me until my attention was 
called to it by Hall's article (Hall 1910). He states that this 
species belongs to the genus Proteocephalus . I quote the par- 
agraphs (Hall 1910: 146-148) in which he gives his reason for this 
belief: "In compiling a key to the dog tapeworms, an examination 
of Kholodkovski' s (1908) description and figures of Taenia punica 
from the dog showed that the cestode in question probably belongs 
in the genus P r o t e o c ephalu s We inland. The head, the uterine 
stem, the position of the ovaries at right angles to the uterine 
stem, and the position of the testes and the genital canals all 
indicate this. The granular strand of uncert: in nature v/hich 
Kholodkovski noted in the position of the excretory canals can 
hardly be anything other than the vitellaria, in the location 
usual for species of the genus Proteocephalus . Kholodkovski states 
that the vitellarium is very small, but it seems likely that he 
has mistaken something else for the vitellarium. A comparison of 
the figures with mounted specimens of worms of the genus 
Proteoceph&ius leaves no reasonable doubt on this point, and it 
is the opinion of the writer and Dr. B. II. Ransom, with vfoom 
the point was discussed, that it is more likely that the dog 


from which the tapeworms were obtained had just eaten the true 


host, some fish, reptile, or batrachian, than the dog was the 
true host of virtue of a normal, even though unusual, infection 
with larval form. Fuhrmann appears to have overlooked the usual 
features of this worm in his review of Kholodkovski (1909), and 
states that the anatomy is that of species of Taenia." - 

" Taenia punica Kholodkovski , 1908, should therefore 
be known as Proteocephalus punicus (Xholodkovski , 1908 Hall, 
1910) a combination proposed here for the first time." 

After studing Cholodkovski 1 s deseript'on and drawings 
and comparing them with the many specimen of Proteocephalidae at 
my disposal I an satisfied that the normal host of this species is 
not the dog, hut a snake. This cestode on account of having its 
testes in two fields does not resemble any of the species of 
Proteocephalus thus far described from fish. The Proteocephalidae 
that infest the lizards belong to the genus Ac ant ho taenia which 
is distinguished fron other Proteocephalids by its spines on the 
head and neck. The Proteocephalids thus far described from 
amphibia are small with rather small heads. Their genital organs 
however resemble those of Taenia punica . The species from snakes 
vary in size from small to large. 


Some oi' the species have large heads with large and prominent suckers 
Their genital organs much resemble those oi Taen ia punica . As in 
Taenia punica the vitellaria oi' the Proteoceohalids oi" snakes are 
sometimes composed^small follicles which rel'use to take the stai . 
well. In such cases it is cas, to overlook the vitellaria. As 
an instance oi' this might be mentioned the Taeni a eunectes A. J. 
Smith in which the coils oi the interovarial space were thought to 
be the vitellaria. For these resons I believe that the true host 
01 Taenia punica is a snake and that Taenia punica belongs to 
the genus Ophiotaenia and should therei'ore be known under the 
name Ophiotaonia punica ( Choiodkovski, 1908.) La due, 1911. 


1855: Ta enia lactea Leidy , 1855:443. 

On account of the inaccessibility of Leidy 1 s description, 
it is here quoted in full: 

TAENIA LACTEA, Leidy. Head small, continuous with the 
neck, without rostellum; acetabula anterior, hemispherical, situated 
at the four angles. Feck moderately long. Segments anteriorly 
transversely oblong, posteriorly longer than the breadth, square 
with rounded angles. Generative apertures marginal (indistinct in 
the specimen. ) 

HAB.- One specimen 16 inches long and 2/3ds of a line 
wide was found in the intestine of T ropidonotus sipedon . In alcohol 
the specimen contracted to one-half the original length and widened 
to 1 line. 

This is probably a species of ~ Qphiotaenia . and may be 
the same as the species reported by me from Nerodia ( Tropidonotus ) 
rhomhif er . Leidy' s description is too inadequate to permit a care- 
ful determination. This form must be included in the list of 
species inquirenda. 



(PA t /<?• /wJ 
l v JU8: Ichthyotaenia p igment at a V. Ifnstow 1908:85. 

von Linstow ( lyOc described this species which Edward 

Jacob son had collected in Java fSemarafig^ Unfortunately the 

specimen was immature so his description is necessarily incomplete. 

This is probably a species of 0phiotaenia « The presence 

ol lateral vitellaria removes it i'r'om the genus o6ch^ r j ati p.a. It 

must be placed in the list ol" incompletely described speces. 

(Taf. 1, 1-ig. 12 und 13). 
Aus der Schlange Psammodynastes p ilven i©ntus Fisch.- 


Lange 32 mm., Breite vorn 0,51 mm., hinten 1,78 mraj 
ein sehr lange strecke ist ungegliedert , nur der letzte 2,6 mm. 
lange theii zeigt deutiiche Proglofcidenbildung; es ist nur ein 
noch nicht vollig entv/ickextec Exemplar vorhanden. DerScolex ist 
abgerundet und nicht oreiter ais die rolgende Strecke; die lange des 
Scolex bexragt 0,57 mm.; die kreisrunden Saugnapfe messen 0,18 mm.; 
ein Postellum und Haken fell en. Die Glieder am Hinterendo sind 
0,13 m. . lang; ihre Breite oetragt 1,1mm., die D:cke 0,25 mm. Die 
Cuticula ist u/:<052 mm. lick und tragt an ihrer Auscenseite einen 
0,o0b5 mm. dicken Stabchenbesatz. Die Muskuiaxur ist stark entv/ickelt 


und man untorscheidet 6 verschiedene muskeliartei;, 1, Ring. und 2, 
Langsrauskeln in dunnerer Lage unfcer der Cuticula; an Parenohymmuskolr 
3. aussere, 4. innero Langsmuekeln, icuztero ocstenen aus Lluskei bund- 
em, t> Dorsoventral- und 6. Transversaimuskeln. In jedem Quersonnitt 
erkannt man fctwa 20 Langsgef asse; die Hauptlangsnervenstamme vcrlaufs<:n 
l/t> des Quer dur chine ssers der Proglottide vom Kande entiernt; masses 
hart I'inden sich dunkei pigment irte t meistcns eiiorraige, durchschni- 
ttlich U,u26 mm. lange und u,u2i mm. Droite Kalkkorperchen. ITur in 
den letzten Progiottiden findet man eine Anlage der Gesohleehtsorgane , 
besonders deutlich ist jederseits nach innen vom Nerv der Rundiiche 
Dotterstock; trotz der geringen Sntwicklung der Gesohleehtsorgane 
ist es zweiioiios dass die Art zu Ich ^hyotaenia gehort!' 


P/> 7, />'?'■ 1 3 » -Wis r/S.frp- /55, tn) 
1350: Taenia racemoaa Diesing, 1850: 511, in part. 

I860: Totrabothrium Gerrardii Baird, 1860:446-48. 


Tetrabotiiriun Gerrar&ii 

Baird, 1861:228-30. 


Tetrabothrium Gerrardii 

Diesing, 13C4: 82. 


Ichthyo taenia Gerrardii 

Luhe, 1898:652. 


Crepidobothriun Gerrardii Monti cell i. 1399:9 


Ichthyo taenia Gerrardii 

Luhe, 1899:525. 

1905: Taenia racenosa Shipely, 19.5:101. 
1908: Taenia eunec tes Smith, 1908:39-41. 

SPECIFIC DIASNOSIS.- Characters of genus. Length of 
strohila as rnuch as 45.6 ens. Max breadth from 0.85 to 2.00 and 
even to 6.0 mm. Strobila flat, thin, ..wrinkled. Proglottids closely 
attached. Segmentaion usually indistinct. Head, lage, tetragonal, 
pyramidal. Mas. breadth of some from 0.935-1.3 or even to 1.75 mm. 
Thickness a little less than breadth. Length about 0.68-0.90 mm. 
Fifth sucker vestigial but with cavity open to extsrior. Four sucker 
prominent, muscular, .Cordate, with lov7er margin interrupted and 
reentrant into sicker cavity. Breadth of suckers 0.40-0.50 mm. 
broad. Heck broad, not over 0.5-0.8 mm. Long, First proglottids 
short, increasing in length rapidly. Mature proglottids quadrate. 
Ripe proglottids longer than broad. Length of some 1.7-2.3 mm. 
long by 0.65-1.09 mm. broad. 


GE1TITA1 -ORGANS.- Genital pore marginal, irregularly 
alternating, situated near middle of proglottid. Vagina anterior 
or posterior to proglottid. Opening of vagina dorsal to cirrus- 

Male organs.- Testes 200-4000 in number, small 0.035 
-0.050-0.080 mm. in diameter, situated in two lateral fields. 
Cirrus-pouch 0.23-0.2,5 mm. long "by 0.10-0.15 mm. "broad. length of 
same 4-5-5- times into proglottid breadth. Several coils of ductus 
e jaculatorius. 

F emale organs .- Lumen of vagina near opening large. 
Sphincter vaginae heavy. Receptaculum seminis present. Ovary 
posterior^hilohed^ lohes thick, short, irregular. Organs, of 
interovarial space as i# Ophiotaeni a . Vitellaria small, loosely 
follicular, lateral. Uterus when fully developed with 20 to 30 
lateral t> ouches. 


J1GGS.- With tw^ or three membranes. Outer membrane 
0.085-0. 100 mm, second membrane 0.028-0.030 mm. , embryo 0.01o-0.018 
mm. in diameter. 


Had ± x • 

Tn intpstine 

f Boidae, a family of South 

American snakes. 

- " 



Collector ; 


?Ophiomorphus Miliaria CO 06b iwlll lllub 

Museum Vienna 

:i if 

Diesing, 1850; 

ti n 

Tl II 

ii it 

X UpillS :.iC i 1 3:7111 

: llatterer 

n n 

Boa constrictor 

Enne c t e smuer inus W agl . 

Siinectos murinus 

:So. America 

: Edward Garrard 

,Baird, 1860:446- 

•(Shipley 1905-101 

La Rue. 
femith 1808:39-4} 
La Rue. 

- A. J. Snith 

1) Spelled mar.racca by Diesinf?. 

In the discussion of the hosts of Ophiotaenia racemosa 

(p.) I otated that I considered that 

Ophiomorphis miliaris and 

Ophis merremii were probable hosts of that species. 

I doubt very 

much if they ever harbor C. Gerrardi" 

.1 consider that Bothrops 

jararaeca which is one of the Crotalidae cannot normally be a host 

of Creoido bothruim gerrardii. but that it would be the host n-r « 

species resembling 0. 1 

larenzelleri. 0. grandis. 0. trimeresuri. 

and 0. Calmettei. These form a erroup distinct f-rnm nth^r 

Ophio taenia. It is very likely that 

Diesing confused several species 

of cestodes under the one name Taenia racemosa. 


The hosts of C. gerrardii of which there is no doubt are 
Eunectes murinus Wagl. and Boa constrictor L. 

HISTORICAL SUMMARY . - This species was first reported 
"by Diesing (1850:511) who called it Taenia racemosa Rud. Besides the 
diagnosis ) which agrees excellently with that of Or e p i d oh o thr ium 
Gerrardii Diesing includes as host one of those species Eunectes 
murinus (scytale) from which this species has "been reported several 
times. I quote his description and hahitat in full: 
^Taenia racemosa RUD01PHI. 
Caput magnum tetragonum, acetahulis angular ibus subter- 
minalibus v. terminalibus , subovatis v. cordatis. Collum nullum. 
Articuii supremi brevissirai , suhsequentes longiores et latiores, 
angulis rotundatis, ultimi longi parallelopipedi angustiores. Penes 
filiformes basi incrassati, marginales vage alterni. longit. 2 # _ • 

l£ rt ; latit. med. 1 3"; ultim. 1". 

Taenia racemosa Rudolphi: Synops. App. 6 C J2. Dujardin: 

Hist. nat. des Helminth. 610. 

Habitaculum. Ophiomorphus miliaris, Martio. Eunectes 

Scytale, Octobri, Fehruario et £unio. Bothrops Mararacca, Martio 

et Aprili. Ophis Merremii, Januario, in Brasilia (batterer): in 

intestinis. M. C. V. ?r 

Ba^/d (1360:446-48) described this species which was found 

"by Mr. Edward Gerrard in the intestine of Boa constrictor from South 
America. Since the name Taenia r acemosa Rud. had already been 
applied to a different species of cestode the name suggested by 


Baird stands by reason of priority. Baird second description 
(1851: £28-30) is a copy of his description (13 60:446-48). On 
account of the inaccessibility of either of these descriptions for 
many investigators I quote the later one in its entirety: 
Tetrabothrium Gerrardii Baird . 
Caput magnum tetragonum, bothriis quatuor cruoiatim oppositis 
subcircularibus magnis, marginihus eontlguis, sing^io costa.... 

r 1 

'°Each of them large, round and having on one side a strong ridge " 3 aire! 
\ 7 

1. c. Os.... Collum nullum. Corpus depressum augustum, articulis 
supremis angustissimis , poster ioribus quadrangularibus. Aperturae 

genitalium Longit. fragmenti 8", latit. capit. 3/4. 

Tetrabothrium Gerrardii Baird ; in Proceed. Zool. Soc. London 
1860. 448 et in Ann. nat. hist. 3 ser. VII. (1861) 230. 

Habitaculum. Boa Constrictor: ex American meridionali, in 
intestinis (Gerrard). 

The genus tetrabothrium was founded by Rudolphi in his 
"Synopsis. Sntosoorum for containing species of Bothriopcephalus 
which possess four bothria. It was afterwards fully adopted and 
characterized as a distinet genus by Diocing; and nine species 
are described by him. These were discovered all inhabiting the 
intestines of certain Mamnalia, B^rds and Marine Fishes: but none of 
them have been, till now, recorded as liTing Reptiles. 

The species now to be described, however was found, by 
; Mr. Ednard Gerrard of the Briish Museum in the intestines of a Boa 
; constrictor from South America. 


The head is large, tetragonal; the four bothria dis- 

I posed crosswise, joined by the margin; each of them large, round and 

having on one side a strong ridge. Body depressed narrow, articulated 

No distinct neck. Anterior extremity of the "body very narrow, and 

the articulations there are extremely small, "becoming larger as they 

descend, the inferior "being quadrangular and rather large. The 

i margins of the articulations somewhat annulated, hut having no ap- 

i pearance of genital apertures. The head is about f of a line broad, 

but I could not discover any mouth. Apparently only fragments of 

these worms here obtained but some of these detached pieces were 

about 18 inches in length. 

Habitat: Intestine of Boa constrictor . 

Diesing (1864:82) placed this species in the genus 

Te trabothrium but with some reservation. I quote his diagnosis in 

in full in order that a comparison of it and his diagnosis of Taenia 

racemosa Diesing (1850:511) can be made: 

Tetrabothrium Gerrardii Baird. Caput magnum tetragonum, 

bothriis ouatuor cruel at im oppositis subcircularibus magnis, margin- 


ibus contiguis, singulo costa.... J Os Gullum nullum. Corpus 

depressum augustum, articulis supremis angustissimis , posterioribus 
: qua&r angular ibus. Aperturae genitalium . . . .Longit. fragmenti 18", 
latit , capit . £ w ' . 

Habitaculum. Boa Constrictor ; ex America meridionali, in 
intestinis (Gerrard). 

A comparison of these diagnoses of Diesing (1850:511) and 

(1864:82) leave no dount that his earlier diagnoses concerned the 

l.("Each of them large, round and having on one side a strong ridge" 
Baird I.e. ) 


sane species as the later one# 

Lfihe (1898:652) included this species in a list of 
Ichthyotaenia from snakes. He gave no reason for considering it a 
member of this genus. 


Fuhrmann [ 1899:864) in a footnote made this statement 
concerning this species: Liihe, M. , Oochoristica nov. gen. 
Taeniadarum. ( Zooio. Anz. 1893. ITo. 576) gjebt an, dass T. Gerrardii 
(Baird) in daB genus Ichthyotaenia gehore; dies ist aber, wie mir 
Prof, fflonticelli raitteilte und wie i"h mieh seibst an den von ihm 
nach den Originalen angef ertigten Zeichniingen uberzengen konnte, 
keineswegs der Fall. Es gehort diese Form in oin besonderes Genus, 
das Prof. Monticelii demnachst unter dem Hamen Grepi'ootnrium 
publizieren wird." 

During the same year Monticelii (1299:9-25 ] made a 
report ac ompanied by drawings, a careful study of the type material 
of this species which was in the British Museum. He also studied 
some material which hsd been sent to him by Prof. fr. Bell. As a 
result o J ' this itudy be erected the genus Crepidobothrium for Baird' s 
specier alone. 

Liihe (1099:525) in a footnote discussed the position 
of this species which he still maintained belonged to the Genus 
Ich t hyot aenia . His ground for this belief was that the head and 
suckers , which are different from those of other Ichthyotaenia, 
constitute only a specific difference. He stated that the name 
Crepibothrjum Puhrmann 1899 ,.as a nomen nudum. He had not seen 
Monticelii' s article and so v/as in no position to judge Monticelii *s 


Shipley (1905:101) made the following report of this 
species , "T aenia raoemosaR ad. Diesing, b yst. Helm. I. p. oil. Many 
specimens from E uneaten murinus Wagl., it has also been described 
from the intestine of B ot "rops j araracca Wagl. " Thanks to Prof. 
H. B. Ward who secured ^for mp- study, I have been able to examine 
some, of Shipley's specimens which had Sunectes murinus as the host. 
They were probably from the same lot which Shipley recorded in the 
above quotation. They proved to belong to the species Cr ep idob othr iu in 
errardii (Baird). 


Smith (1908) found some in an anacoda, Eunectes 
murinus . He described them under the name T aenia eunectes . 
Doctor Smith sent some of this material to Doctor Ward who has very 
kindly placed it at my disposal. Study of the material has shown 
it to be Grepidobothrium gerrardii . 

DISCU3SI01I.- A comparison of the diagnosis oi 
Diesing (1850:511 and 1864:82) with the description by Bain 
and with the description by Mont i cel. '.i (1899:9-25) leaves no ground 
for doubt that these investigators were reporting the same specjes. 
In certain respects the description by SJaith (1908:39-41) differs 
radically from that of Honticelli. However an examination of Smith's 
material has shown that Smith made numerous misinterpretations which 
are the cause of the apparent 


discrepancies between his work and Monticelli ' s. In my study of 

this form I have found Monticelli correct in the main. It has alread r 

been stated that Shipley's material "belonged to C. G errardii . 

The question concerning the norrect name for this species 
must be settled. Prom a structural point of view tnifl species 
cannot he retained in the genus jFaenia nor in the genus Tetrabothrium 
Anatomically this species is one of the Proteoce phalidae . 

This has been recognized by iiuhe (1898), Fuhrmann (1399), 
and Monticelli (1899). The name which Fuhrmann (1899:364) gave for 
it can be dismis ed as a nomen nudum. Luhe 01898 and 1899) 
considered that the species belonged to Ichthyo taenia . I have alread , p 

shown in the discussion of the genus Proteoceph a lus that the name 
Ichthyo t aenia Lonnberg (1894) is a s~;non3:m of the name j- -otoocephalue 
Weinland (1858). 

The latter name also antedates the name Crepidobothrium 
Monticelli (1899) and on that account should be used in place of 
Crepidobothriura if structually Baird's species agrees with Taenia 
f ilicolli. RiaAz a mbigua , the type of Proteoce phalus . An examination 
of Llonticollis (1899) description and figures and of ray description 
with its accompanying figures, shows conclusively that this species 
does not agree anatomically with the type of Proteoce phalus and 
that this species really belongs to a flifferent genus. The only 
avialable naraeis Crepidobothriurn Monticelli, 1899. 


Source of Material.- One bottle if 9 specimens with 

heads, labelled Ichthyotaenia g crrardii (Baird)?. aus Ertnectes 
murinas intestinum Berlin Aquariei»i was received from Dr. Anton Colli:. 

Berlin in answer to a request from Doctor Y/ard. This is now Ho. 
179 in Prof es.. oi Ward's collection. 

One bottle of 5 specimens with heads and some pieces 


"Taenia racemosa Rud. from Eunectes murinas." was receive 
from A. E. Shipley, Cambridge. The materia/ is now 1T0. 08.472 in 
Professor Ward's collection 


One bottle of 3 specimens with heads labelled Taenia 
Eunectes from Anacorda.$m. Intestine. Phila. Zoo. G. 939. U. ?. Path , 

Hist. 1694 was received from A. J. Smith, Phila. This is now Ho. 10.19C 

in Professor Ward's collection. Seven slides, one of them containing 

w , 
a head labelled, ITo. 1858, From Boa bonstrictor Box 53B0, 1823" were 

received for examination from the Bureau of Animal industry. Washing- 


COMPARATIVE STUDY OF MATERIAL.- A careful study of the 
Specimens thus "brought together showed them to "belong to the same 
species. In all cases one or more heads were present ao that a 
careful comparison of them could be made. Some toto preparation 
were made from specimens of the alcoholics. WM e other were cleared 
and examined in glycerine. On acoouHt of the small amount of materia 
the section method was not used except in a single case. The table 
(p. ) best shows the comparison of the scolices, suckers and necks 
of the various lots, The heads used for measurement were chosen 
quite at randon for an examination of the material showed that a 
tabulation of measurement of all heads to be unnecessary. 

Comparative table showing dimension of heads, sucfeers and 
necks of C. Gerrardii from various sources til measurements are in 

Lot number 







Sucker s / yirr s e; 

diameter -0.646 


breadth a. 000 
length £.500 1 

unf l;-.tton 

0.680 ± 





OS {« 2f rorflO . 1 79 : 1U -179 ; 1Q #1 
Snipxey in"Anton, ; Anton :" AJ# 


0.680 ± 

. .646 


Collinin .Collin :smith 
g] ye em n q " J n ^ y < i n S3 y 

.10.179 rio.190 -13?*^ 


1.190 : 1.090 :1.190 

0.680 ± : 0.680 i0.850 

0.544 ]0. 510 -0.510 

0.544 . ' : : 0.544 

0.402 -0.390 : 1.190 

0.850 ■ 0.500 0.500 



0. soq 

1. Eoqb 


(*) Head showed a little effect of flattening in mounting 
This may have had some effect on suckers also. 

("b) Head evidently very strongly contracted as were also 
the suckers. 

(c) Measurements were taken from Smith's drawing;. which 
was accompanied by a scale of magnification. 

A study of the above table shows that in the main the 
measurements agree very well. V/here apparent discrepancies occur 
they are easily accounted for by states of contraction end com- 
pression. In every ca^e the shape of the suckers is the same, due 
allowance "being made for contraction and relaxation. Every haed 

of the form lots was examined in this respect. In each case the 


lower margin of the suckers^ turned in the an angular intrusion into 
the cavity of the sucker as Smith (l c 08) described it. In the more 
contracted or in greatly expanded suckers this condition was somewhat 
difficult to make out but -./hen properly cleared as in glycerine or 
balsam it was apparent. A study of toto preparation of the proglottid 
from the various toto fails to disclose any distinguishing 
characteristics between the different lots. I shall ther fore "base 
the following description upon this material so collected together, 
referring 1at times to special lots. I shall uso the descriptions by 
Smith (1908) and by Honticelli (1899) as sources of comparative data. 
Jet was desired to study the slides upon which Monticelli based his 
jLescription but unfortunately Professor Ward was unable to secure 


EXTERNA! CHARACTERS.- This is one of the largest of the 
Protebcephalid species inhabiting snakes. 0. Gal e ttei Barrois, 
0. Uarenjtfglsel l eri Parona, 0. P randis are of ah out the same size. 
The latter even exceeds P. Gerrardi i in some respects. ITe^t smaller 
than this group comes 0. racemosus (Rud.). A comparison of the 
table at the end of this report shows the size relationships of these 

The strobila varies considerably in length. According 
to Baird (I860) it measured 45.6 cms (18 in diet. Smith (19 8) 
reports specimens 95-100 mm long. Diesing (1850:511) gives its 
length as 2 inches to 1' feet. IJonticelli gives no data on this 
point. The reported breadth varies from 0.85 mm in some short 
specimens eKar.ined by me to 1.8 mm in the B. A. I. slide 1858, to 
2.0 mm, according to Smith, and from 2*5-6.0 mm (1-3 lines) in the 
middle to 2 mm (1 line) at the posterior end, according to Diesing 
(1850). The strobila is flat and relatively thin. The surface is 

j more or less thrown into folds. The proglottids are attached by 
their full width. The segmentation is indistinct in the anterior 

, region and more distinct in the region of mature and ripe 
proglottid;:. In certain strobila the segmentation is more distinct 
than in Gthers. It is never as clearly marked as in some species 
Proteocephalis. . 

The head is large, readily distinguishable to the naked 
eye. Baird describes it in there words, "The head is large tetragonal 


the four bothria disposed arosswise, joined by the margins. "Smith 
A.J. -(1908) says of it, "The head, viewed from the front presents 
a conical appearance from the prominence of the suckers, measuring 
tansversely across the two opposed suckers 1.5 mm. and laterally 

across two adjacent suckers 1.3 mm. The suckers, thus prominent, 

form the rounded arms of the crucial frontal picture, each sucker 
being globose in shape and having a lateral diameter of about 0.7 

.mm. " 

Xonticelli's (1099) description of the head reads: 
Come si vede, il capo^IJonticelli ' s figure is reproduced [PI. fig. ) 
e alsrbastanza gran&e e ben distinto dal collo che anteriormente si 
slarga di poco come pei aostenerlo; esso ha Vaspetto grossolano 
di an porno da bast one e, come si rileva dalla figura, e rigonfio e 
massiccio, tetragonale, a forma di piramide tronca. Anteriorr.ente 
ristretto, cupuliforme, terminate nel mezzo da un indistinto 
cocuzsoletto apicale conoide; posteriromcnte slargato e troncato 
sparge oltre il collo pc i suoi spessi margini rigonfi e presenta, 
nel mezzo di ciascune dei ouatro lati, una insenatura molto accentuate." 

The head is always tetragonal and somewhat pyramidal. 
These shapes are varied to some extent by the states of contraction. 
A head which is relaxed and whose suckers are expanded may have a 
lobate appearance. Such a head is figured (PI. 7 f ig#//^) where 
deep grooves may be seen between the suckers. In greatly contacted 
heads the suckers are drawA closely together, the furrows between 
the suckers are almost obliterated and the suckers themselves are 


reduced in size. Such a head is figured (PI. 7 fig. 146) in which 
the suckers are seen to be forward while in the relaxed head the 
suckers are directed outward. These figures shoiild be compared with 
the drawings of heads by Smith (1908] and Monticelli (1899), which 
have been reproduced here (PI. 7 figs. 138-139 ) and (PI. 8 figs. 
155-156). These figures illustrate pretty well the various states 
of contraction which one meets in a study of considerable material* 

The dimensions of the head are; breadth 0.935-1,30 in my 
material, 1.75 mm. (Baird), thickness nearly equals breadth, length 
about 0.68-0.90 mm. 

The apex of the head is slightly elevated, rarely flat. 
There is no rostellum no spines. 

In toto preparations I could find no trace of a fifth 
sucker but in a series of transverse sections I found a small sunken 
, fifth sucker which had degenerated until it had lost its raial 
musculature. The sucker still possessed an opening to the exterior 
measuring 0.037 by 0.026 mm. The cavity was but a little more than 
0.030 mm deep. The sucker measured about 0.048 x 0.067 mm on its 
transverse axes. From the tip it extended about 0.060 mm into the 
head. While the radial musculature could not be demonstrated in 
my preparation the basement membrane was still well developed. The 
tissue of the sucker is composed of rather large elongated cells, 
which plainly no longer functions as muscle cell. 


This sucker ..-sands about mid-way between the vestigial sucK r 
Of 0. filaroides and 0. Lonn'oergii and P. amblophi bis on the one 
hand and the functional fifth sucker of P. fall ax P. -oinguis and 
p_. percae on the other, , 

Approximately the same stage of degeneration i: shown 
in PI. fig. which pcrtrays a degenerating fifth sucker of the 
pleroceucus of 0. f ilaroides . 

Suckers .- Or these organs Monticelli (10 c jyJ writes: 
"Hon vi sono botridii, ma ouattro ventoso, grnadi ovoidali, disposte 
in croee, eecondo il loro asce maggiore, ai qu£.ttro angoli del capo 
ed obliguanentc e convergent! verso il oocuzzoletto apicale. insi.e sonf 
approfondate nello spessore del caop e ne occupano quasi tuta la 
lunghez^a; fra le ventose il caop si iniossa in piccoli solchi long- 
itudinal che si terminano nelle imsenature marginali innanzi ricordat 
Le ventose hanno forma di ferre di cavalo e, diro meglio, ricordani 
la figura di un pie do di cavallo ferrato visto di sotto. Es. e hanno 
un aspetto carateristico proprio, ehe puo facilmente ricavarsi dalla 
fig. 2 e, meglio ancora, dalla fig. e, che mostra il caop visto 
di sopra ed alquanto schiacciato. Che il margina posteriore delle 
ventose non e integroa ma scavato do una insenatura formata dal 
ripiegcrsi verso il mezzo del cavo di esse delle oareti posteriori, 
della ventose — che non formano un tutto continu , ma sono posterior] 
mente intorrot'.e — che ravvicinate fra loro, costituiscono una 
doppia crosta, breve, la quale si perde, divaricando in due rametti 
a V, nel fondo della ventosa. Ma, meglio che dalla descrizione 


puo interdersi la peculiare struttura di queste vontose, gia admom- 
hrata.dal 3arid, come si rilevza dalle perole innanzi citate, dall' 
esame delle figure z e 3, che danno una fedele immagine della forma 
del capo del cestode in esame, come I'ho ricavata dallo studio del 
tipe origin- le del Barid." 

Monti eel Hi 1 s figures above mentioned are reproduced (PI. $ 
& igs In these figures the point of the intnrned margin 
is split and each arm of the"V." is turned out. My material shows 

no such spreding of the point of the "V". : A series of trans- 

sections how that the point of the "V" is closed. When the sucker 
is much relaxed the point is difficult to see, but it may be found 
if the head be cleared and examined in glycerine, likewise in very 
strongly contracted suckers the inturned point may escape detection 
except when cleared and examined under favorable conditions. From 
my examination of three heads from Smith's mat .rial I am letf to 

"believe that; zhq narrov.- slit-like opening in the suckers of his 

s~ ^ i l3 *>' 3f 

figures^are due toN ^reprnduoed PI . 7 figi^ foreshortening. Usually 

the opening is much more nearly round than here shown. 

The suckers are very prominent and are most prominent in 

a relaxed head (?1.7 fig.//£). They measure 0.40-0.70 mm. in 

"breadth. On the average they measure ah out 0.55 mm. 

ITeck .- A poorly defined neck region separates the head 
from the body _ Barid (1860) says:" Ho distinct neck. Anterior 
of the body very narrow". 



Monticelii (1899) says: II collo ' e relative amenta lunghotto e la 

prime proglottid sono come delle rughe e molto indistinte". 

for the neck his 
A. J. Smith (1908) gives a width of 0.300 mm. and found 

no- drawing a length of 1.20 mm. llj own measurements of Smithes 
material however gives width up to 1.190 mm. Hy measurements of 
6 atrobila give a length of neck of 0.500-0.850 mm and in these cases 
it is probable that the measurements given ars too great. The region 
of proglottid formation begins very close to the head and here the 
proglottid:? are very narrow anu their boundaries poorly defined. The 
neck is thin and flat. 

The first proglottids are very much broader than long. 
Ag they become older they increase rapidly in lenght. Mature segments 
are nearl" quadrate while ripe proglottids are longer than broad. 
The moniliform proglottids mentioned by Smith 1908) are not charactr- 
istic of the species tut are due to contraction states. Among the 
three specimens which he sent to Professor Y/ard a short one had 
these peculiarly contracted proglottids. The longest and most perfec 
specimens had no segments of this charcter nor were such proglottids 
observed among she specimens of the other lots. Ripe proglottids 
measure 1.70-2.30 mm. in length by 0.65-1.09 in breadth. 

IUTBRUAI ANATOIIY. - The genital anlagen appear very early 
in the chain . In fact they are to be seen in all but the youngest 
proglottids. Tho genital aperture is irregularly alternating 



It is situated near the middle of the margin of the segment either, 
anterior or posterior. The vagina and cirrus open into the common 
genital sinus, the vagina being either anterior or posterior to the 
cirrus-pouch. A genital sinus can scarely be said to exist. Under 
favorable circumstances the sinus may be seen to be 0.020-0.030 
mm. deep. This shallow sinus is eafily overlooked. 

The opening of the vagina is sometimes dorsal to the 


cirrus-pouch. I am not prepared to say that it ar.vays A that position- 
Monticell (1899) thinks that the vagina normally lies above the 
cirrus-pouch and that when i:. is found anterior or dorsal to the 
same it has been displaced by pressure. From my work on this 
species and also several other species in which the vagina may lies 
in either the mterior or posterior position. Iam sure that pressure 
has nothing to do with this variation. My preparations have all been 
made without compression. Yet the vaginae alternate irregular 1 
in position. Sections in a frontal plane thrt*. developing proglottids 
of 0. L onnaergii show that the vaginae, sometimes anterior and 
sometimes posterior to the cirrus- ouch. They devlop in those 
| positions which they hold in mature and ripe proglottids. 

A careful examination of toto preparations o~ se7 rel 
species of O phio ta e nia and also o O re p i dob othr ium fle r r ar di i shows 
|that the vagina if in the anterior position extends farther anteriad 
lefore bonding towaed the genital pore than if it lies in the 
iosterior position. If the vaginae are displaced by pressure then 
;hey ou^ht to give evidence of such displacement by thtf er appearance. 

i l 


This they do not do as an examination of my drawings (Pi. 7 f igs«^jL 

will "show. It will be noted that when the vagina is anterior 

to the cirrus-pouch is usually crosses the coils of vas deferens "but 

when posterior it does not cross the vas deferens* In O. G-errardii 
the va~ina when anterior never crosses the cirrus-pouch. 

Llal e organs .- The testes lie in tv/o broad fields which 
tend to coalesce at the anterior end of the proglottid. They number 
about £40 in the specimens from the Bureau of Animal In&udtry. 
Monticel_i (1899) found about 200 or more in each proglottid. Smith 
(1908) figures but 87 testes. There must be an enormous variation 
in the number of testes or else Smith overlooked some of them, 
for in a preparation of an immature proglottid of one of his 
spefipmens I counted nearly 400. This proglottid is delineated 
(PI. 7 fig./^f). In this proglottid, the testes were very small. 
In maturity many of them would probably be crowded down to a lower 
plane and so would be overlooked. I doubt if any of the anlagen of 
testes fuse together or degenerate. As a rule the testes measure 
from 0.030-0.050 mm in diameter but in a specimen from lot 10.179 
the testes measured as much as 0.080 mm. Llontieelli (1399) says they 
are about twice the size of the vitelline follicles. 

The vas deferens ( PI. J flgjfi) in ripe proglotLids 
is thrown into numerous heavy coils filled with sper :atozoa. The 
coils extend nearly to the middle of the proglotttic . The ductus 
ejaculatorius makes a few coils in the cirrus-pouch. The cirrus 


itself , when protruded, is short, thick and very muscular. 
Uonticelli 's description agrees with this statement. 

The cirrus-pouch is short, "broad and heavily muscled. Its 
length is about 0.23-0.H5 mm and its breadth about O.IO-U.15 mm. 

In maturing proglottids the ratio of its length to the segment 
breadth is 1:5 or 1:6 while in ripe proglottids it is about 1:4. 

Female organ s.- These organs (PI.? figs.^ ) are arranged 
as in the O phio taenia . The lumen of the vagina near it opening to 
the exterior is la-ge. The vagina is here surrounded by a strong 
sphincter vaginae. There are no coils of vagina anterior to the 
ovary. I have not seen a receptaculum seminis. Uonticelli (1399) 
states that it is present. 

The ovary is bilobed and is situated in the posterior 
part of the proglottids. The lobes are thick, broad and shcCrt. 
Their outlines are more or less irregular. 

An o6'capt, ootype, shell gland, oviduct, unpaged 
vitelline duet, lower vagina, and a uterine passage are present and 
these organs are arranged in the manner common to the Proteooephalids 

(Pl.(o fig./lj and Pl.£ £lgj27) .Hontaeelli (1899) gives a long and 
detailed description of the arrangement of these organs upon 
which no comments are necessary. Smith (1908) ,- mistook these organs 
o: the inter-ovarial space for vitellaria. 

The vitellaria are lateral loosely follicul<lY* masses 
which extend thru - put the length of the proglottid: The follicles 
are small, and in some of my preparations they stained poorly. 


It is probable that Smith had the came difficulty in staining the 
vitfillaria and for that reason he overlooked tram entirely. 

" states that the vitellaria lie outside of the 
inner longitudinal muscle layer. I have examined nothing but toto 
preparations which perhaps are not trustworthy for the determination 
of this point, nevertheless I believe Monticelli has made a 
misinterpretation in this statement. 

The uterus (Pl.'7 fig5, A ) is a median tube which in ripe 
proglottids has from 20 to 30 lateral pouches on either side. As 
these pouches fill with eggs they become rounded and the septa become 
indistinct. The uterus is not connected v/ith the lobes of the 
ovary as Smith (1908) figures it. Smith's figure does not correctly 
portray the uterus of this epee'es . The pouches are nore numerous 
and nore closely applied to each, other and less slender than he 
shows them. 

I have not seen any uterine pores. As to the method 
of discharging the eggs Monticelli (1399) writes: "Helle ultime 
proglottidi 1'utcro, pregno e rimpinzato d'uova, sporge alquaiito, 
facendo ernia contro le paret£, sulla faccia ventrale^elle proglottid, 
lion he escervato orifizio estorno permanente/dell 'utero , come quelle 
che Kraemer ha visto in elcuni T etr-. cotylus (p. e. S. f illcollis / ) 
e neppure he constatato de terminal Si uno sbocco prowisorio por dar 

uscita alle uova: pens©, forse, l'uscita delle nova all 'esters 
awenga, come in altpyjas^ per deiscenza. " 1) Kraemer ( 1892:^1. 20 
fig. 42) 


E^GS.- According to Smith (1908) fcae six hooked embryo 
measures 0. 016-0. 018 mm. in diameter, the t./o,, 0.028-0 .030 mm. and 
0.005-0.100 mm. respectively. I was unable to measure or to study 
any eggs of this secies hence I am do not feel justified in 
commenting on Monticellis description of the egg. His description 
f oil or; s : 

"Le uova uterine, relativamente piccole, hanno un guscio spesso e 
molte evidente: contengono gia l'oncosfera involta in una ve$te, od 
invoglio, die come una teca a netto contorno e non molte spessa, 
per quant© ben distir"^. e^rconda, come in altri teniadi, I'embrione. 
Giascvn uovo, con il relc/.ivo gascio, e racciuso in una cqpsula grand 
sferoidale e di diametro assai piu del doppio del guscio, omologa 
analoga, a quella che ei osserva in altri cestodi (capsule uterine, 
od ovariche Auct.). La quale, come altrove ho dimostrato 
Uon~icolli ffr. Sav. . — Sulla conidetta subcuticola dei Cestodi, in: 

Rend, R. Acc. SC. ITapoli, Fasc. 7-12, luglio-de<*embre 1892 ( sul princi 
dovo parle del. 'uovo dei cestodi e ne riassumo lo sviluppcj nulla 
ha da vedore col guscio delle uova essendo essa una formazione 
molte diYerso , : econdaria e posteriore: circa x'origine di questa, 
oltre quanto ho espresso nel citato nio lavoro, veggasi pure il 
Dianare ^ Diamare V. — II genere Dipyiidium. in: Atti R. Accad. 
Sc. Hap. (2) Vol, VI. II. 7, 31 pp. 3 TavolaJ. Capsula questa che 
ritenge possa e bba venir interpetrata come una sorta di cocon 
e corrispondente a formazioni del genere che si osservano nelle 
uova di altri aniraali e ne e'^condona il guscio. " 



since his specimens cone from a host 
DI3CUSSI01T 0? TAENIA EU1ECTES SMITH.- Since^.hich is 

not uncommon in American Zoological Gar dens and since his paper 

will probably "be widely referred to by American investigators 

I call more specific, attention to certain misinterpretations 

in the description and delineation of this species by A. J. Smith 

(1908) . 

His fig. 4 PI. Ill . which I have not had reproduced, shows 
very clearly that he considerd that form a species of Tannia. His 
attempt to make the organs of this species agree .v/ith those of Taenia 
has resulted in considerabl confusion. The organ which he has labelled 
the vagina is the cirrus-pouch and viee-versa. The receptaculum 
seminis of his labelling is the mass of coils of the vas deferens. 
The lateral vitellaria he has omitted entirely. That mass of coils 
which he labelled vitellaria is the organs oi the interovarial 
space. The ovary in this species is never connected with the uterus 
as he figures it but always thru the single oviduct, ootype, and 
uterine passage as I have figured (PI. 6 f ig./# ) Tae lateral pouches 
of the uterus, I have never seen in this sepcies or in a>»y species 
of Ophi taenia as he figur s them. The uterus which he figures 
i& characteristic of Taenia n ot of a Proteocephalid. 

Unfortunately I have not been able to examine any 
proglottids of Smith's specimens which were in the same degree of 
ievelopment as that sho..n in hi: figure. I have, however, examined 
younger proglottids and these in every case and in every essential 
>articulQ>~ agreed with the specimens received from others sources. 


There can "be no doubt that Smith's specimens belong to Grepido- 
bothrlnm Gerrarflii . Hence his name Taenia e unectes is a synonym 
of the former and should be suppressed. 

RELATIONSHIPS.- This ppecies alor.e belongs to the 
genus C t o p i do b o rhr iua . It finds its nearest congeners in the 
larger Ophio taenia . While in the structure of the proglottids 
and in the arrangement of the genital organs this species agrees 
almost perfectly with the Ophio t aonia there remain two characters ; 
which I deem of sufficient value to warrant a separation of the 
snake Proteocephalids into two genera. These characters arc the 
structure of the suckers and the length of the neck. The first 
character has already been discussed. All the Ophio taenia known 
at present have relatively long necks, i. e. 2-8 mm. long while 
6« Oerrardii -the only species of the genus. 'jo which it belongs 
has a short neck. 

I consider that Mont ic ell i v;as ar.ply justified in 
separating thin species from hi$- genus Tetracptylus sensu 
latu. However since I have shown that his type species of T etraftco tfr. 

lus should be separated from the genus Proteocoohalus and 
even from the family P roteocephalidae I shal] not repeat his argunen - 
in justification of his action. 

Llonticellis (1899) diagnosis of the genus Crepidobothrium 

reads : 


Capo inermc, grande, rigonfio, paramidato: ventose 
ellittiche, posteriormente a margine interrotto e rientrante nel 


cavo della ventosa. Aperture genitali marginali, irrggolarmente al- 
ternant i. Orifizio dell'antro genital e subventrale, largo: 
abocco della vagina dorsalmente e disotto alia sasca del pene: 
ovaric piccolc: testicoli pieeolissiml, numerosi, collecati nel 
mezzfl della proglottide, internamente alia musculatura longitudinal e 
interna, in unica serie. Utero allungato, tubolare, saccixorme. 
Vitellogeni piccoli, numerosi, periferici. Uova ravvolte in una 
capsula gelatinosa." 

His diagnosis does not require amplification but should 
"be restated thus' Crepidobothrium Monticelli. Head large, swollen, 
pyramidal, tetragonal, unarmed^ fifth sucker vestigial. Four suckers 
inversely cordate, posterior margin interrupted and re-entrant into 
sucker cavi.'.y. Genital aperture marginal, irregularly alternating 
Vaginal opening dorsal to cirrus-pouch. Testes small, very numerous 
in a single layer, situated in two lateral fields. Ovary bi obed, 
lob s irregular, relatively small, 

Uterus tubular in mic -field, in ripe proglottids with 
numerous lateral outpocketings. Vitellaria lateral, small, numerous. 
All genital organs within inner longitudinal musculature except por- 
tion of cirrus-pouch and vagina. 3&gs small, with three (?) 
membranes, outer one gelatinous. Type o f genus .- Crepidobothrium 
Gerra rdii (Baird), from Boa c onstrictor . type host; So. America 
(BraziK?), type locality. 




1906: Ichth yotaenia crypt obothiujT Von Ll-nstow, 1906:185. 

Under the name of Icnthyot aenia c rypt cbo.thr ium von Linstow 
described a cestode from the intestine of a tree snake, Chrysopelea 
o rnats Russell, Kurunegala. 

A careful examination of Lis description and figures show 
that it is not a pJT ^ rooeorcphalus species at all hut is probably a 
species cf Oochoristica . The position of the vitellaria in 
PCrot eocephalus is without exception lateral, the vitellaria ex- 
tending nearly the full length of the proglottid. In von Linstow 1 s 
species the vitellaria are globular organs lying just posterior to 
to the ovary. This is exactly the relation that is present in 
Oochoristica Luke . The invaginated or infolded condition of the 
head is probably not of generic importance and nay have no specific 
value. I therefore suggest for it the species Oochoristica 
crypt obethriyBm (V. Linstow) La Rue. 



(pi. H , tl&t.d5f0fr,Sfj f/J 

1891: Tetraootylus coryphiceohala Monticelli, 1891:151-174. 
1894: Iohthyotaenia G o r yp h i g e o h al a Lonnborg, 1394:803. 
1896: Iohthyotaenia coryphicephala Rig'-enbach, 1895:267. 
1900: Iohthyot a enia c o r y o h i o e phal a Brown, 1900:1680-31. 

SPECIFIC DIAGNOSIS.- Characters of genus. Head dilated 
somewhat triangular or globose, when seen from the front, swollen, 
sub - quadrat e , anteriorly elongated to form a conical elevation, 
unarmed and not very prominent, distinct from neck. Breadth of 
head perhaps 0.35 mm. Suckers four, faily prominent, oval in shape, 
about 0.15 mm. long, situated at four angles of head. So fifth 
sucker. Heck very long, narrow. 

First proglottids short, middle ones almost rectangular, 
posterior and last ones decidedly rectangular, swollen in the 
middle. Ripe proglottids about 1.80-H.0 mm. broad by 0.C8-1.0 mm. 
long by about 0.34 mm. thick. Proglottids attached by full width. 
Intersegmental furrows shallow. Corners of proglottids not marked. 
Segmentation indistinct. 

GENITAL ORGASS.- Genital organs as in genus. Genital 
pore marginal, irregularly alternating in first one fourth or 
one-fifth of proglottid length. ITo genital papilla. 

Hale organs. - Testes spheroidal, 0.05-0.085 in max. 
dimension, numerous, 100 or more, situated in single dorsal field 
outside of inner longitudinal muscle sheathe. Vas deferens a mass 
of coils inside the inner muscle sheath. 


Cirrus-pouch 0.286-0.315 mm. long by 0.094-0.105 mm. "broad. Length 

o±* cirrus-pouch 5 to 5 to 6-y times into proglottid breadth. Ductus 

e jEculatorius^numerous coils. Cirrus slender unarmed. 

Female organs .- Vagina and vaginal opening always anter- 
ior to cirrus-pouch. Vagina never crossing latter. Sphincter 
vaginae and receptaculum seminis present. Vagina within inner 
muscle sheathe. Vitellaria lateral, extensive jVentral^n single 
layer in cortical layer. Ovary posterior, bilobed, 

aiigmented by large protuberances, partially outside Of inner muscles 
sheathe. Organs oi' interovarial space as in Proteocephalidae . 
Uterus with many long slender lateral outpoclcetings , ventral, and 
outside of inner muscle sheathe. 

EGGS. - Membranes not observed. Embryos 0.015-0.017 
mm. in diameter. 

YPE.- Slides in collection of Monticelli, from 
specimen no. 571 in the Zoological Museum at Vienna. 

HISTORICAL SUMMARY. - This species was first described 
from Silurus sp. by Monticelli (1891:151-174). For this and other 
nineteen species oi' cestodesfrom fish he proposed the genus 
Tetracotr'lus . without, however, naming a type species. That genus 
was based cn Taenia c o ryph i c e phal a . Lbnnberg (1894:803) listed 
this as one of the species of Ichthyo taenia without entering into 
a discussion of its anatomy. Riggenbach (1396:267) listed this a., 
a species of Ichthyo taenia . Brdttti (1900:1680-81) states that 
.Ionticelli 1 s genus Tetracotylus is based on the 


rather than sub- triangular . The suckers could not he measured 

except in length which was about 0,150 mm. Monticelli's drawing 

of the A (Pl. i'ig. ) shows that the suckers are irregularly 

oval in shape, the longitudinal axis of the suckers are being the 

• The neck in this slide was too badly crushed to permit, 
accurate measuring. It was quite long^upwards of 3 or 4 mm. y even 
then no traces of segmentation could be observed but the neck was 
cut off at this point. 

No young proglottides were to he found in the slides 
examined by me. Pour representative ripe proglottids were meas- 
ured. Their dimensions were l.ys mm. broad by 0.68 mm. long; l:85x 
0.75 mm., 2.0 x 0.7b mm., 1.00 x 1.0 mm. Transsection of ripe 
proglottids are about 0.34 mm. thick. 

The proglottids are attached by their full width. The 
angles of the segments can scarely be seen. The intersegmental 
furrows are very shallow. These observations were" made on a 
limited amount of material so it is possible that some of these 
statements just made would require modification %± considerable 
good material could be examined. 

IITT:-:RiTAL AffftTOHY . Genital organs . The genital organs 
are arranged as figured (PI. ^ 1 ig$f/,$f **^(Pf these, figure 

hat been reproduced from Monticelli (ISUl) and figures tttJl 

_ are original drawings made from Monticelli's type slides 

The genital pore is marginal, irregularly alternating 


and situated at the end of the first one-fifth to One-fourth of 
the proglottid. There is no genital papilla. The genital pore is 
the opening into a small genital atrium into whicn both cirrus- 
pouch and vagina cpen. The vagina always lies anterior to the 
cinrus-pouch and its opening is also clearly anterior to the open- 
ing of tho male ducts. Monticelli notes a sort of sphincter muscle 
about the genital atrium (Pl.f fig-tf>"). Ho states that this sphincter 
draws the opening of the vagina and cirrus-pouch very close together. 

Male organs .- The testes are spheroidal. They measure 
from 0.05 up to 0.05-0.07-0.085 mm. They number ahout 100 or 

even more and they are situated dorsally in a field which covers the 
entire proglottid except that region occupied by the ovary and by 
a narrow strip on either lateral margin of the segment. There is 


no free median zone^Monticelli states and as he shows in his figure 
(' .PI. *f fig. ftf ) which is misleading, His statement that they 
are not numerous is also incorrect. The testes fPl. f f ig. ff ) are 
outside of the inner longitudinal muscle layer, a condition which 
does not occup in any species of the Proteoephalidae not, so far 
as I can aecertain, in any family of EetrapH^llid^lg/. A careful 
study of Brattn's (1004-1D0O) great monograph on the cestodes gives 
no reason for "believing that this condition has "been observed here- 
to fore in any o rder of cestodes. In the P s eud o phyl 1 i d e a he states 
definitely that the testes occur in the Medullary layer. He makes 
the same statement for the T e t rap; :yil ide a and the CyclOwhyllideo . 
Audi as for the genital organs in the Diphyllidea and the 
Trypanorhyncha he states that they are as in the T e t r aphyl 1 i d e a . 


The situation of the testes in this species then is certainly unique. 
The vasa ef f erentia have not "been observed. The vas 


deferens fffl»a*an elongated mass of coils which extend iron the 
cirrus-pouch toward, but not reaching, the mid-field. This mass 
of coils lies just a little posterior to the cirrus-pouch and with- 
in the layer of inner longitudinal muscles. This duct is not of 
uniform diameter thruout but is narrow in its beginning region and 
it is again constructed for a short part of its length before enter- 
ing the cirrus-pouch. The vas deferens with its swollen coils 
functions as a vesicula seminalis. 

The cirrus-pouch (Pl.V fig.^J) is somewhat muscular. 
The muscles are best developed near its outer and. Its length varies 
from 0.286 to 0.315 and even to 0.340 nm. and its breadth from 0.0U4 
mm. in the longer pouches to 0.10b in the shorter ones. The pouch 
(Pl.V f ig. 1/ ) tapers frequently to a blunt TOint at the inner end 
and is usually broadest ne; r its opening. Its length goes from 
5 to 6 or 6 1 - times into the proglottid breadth. 

The ductus ejaculatoriW (Pl.^ figs. #5) is slender and 
has thin walls. It makes numerous intricate coils before passing- 
over into the cirrus. The protruded cirrus has not been seen by 
Monticelli or bgr me. As it appears in the cirrus-pouch it is not 
very muscular, not large, and is not armed. Probably when evaginated 
it would bo long, slender, and without a th ck base. 

Female/organs .- As seen in a toto preparation these 
appear to be anranged in a general as in the Proteoce^halidce but 
a careful comparison of this species with any 

Proteocephalid revels stricking differences which will be considered 
in their proper place. 

The vagina (Pl.y fig.?/ ) always lies anterior to the 
cirrus-pouch and it never crosses the latter nor does it cross the 
lass oi coils of va$ deferens except in rate cases and then it 
Grosses ventrally only the most anterior coils. Monticelli states 
that is crosses the coils of vas deferens dorsally. The opening 
of the vagina is always clearly anterior to the opening of the 
cirrus-pouch. In many bpecies of Proteocephalus and also of 
3phic tg.enia the opening is dorsal to the cirrus-pouch and I am incl- 


ined to believe that if the renaining species of those genera were 
carefully investigated with that point in view they too would show 
the opening of the vagina dorsal to the cirrus-pouch. Near the 
vaginal opening there is a weak sphincter vagina. Beyond the 
sphincter the lumen of the vagina dile.tes somewhat for a distance, 
perhaps 0.3 mm., then it is constricted. After arriving in the 
cid-field of the proglottid the vagina dilates again considerably. 
This dilatation persists thru several curves and coils of the 
vagina until the latter is just about to enter the interovarial 
space then the vagina is greatly constricted. The dilated region 
is the receptaculum seminis which is much more merked than in any 
species of Proteooeohalus observed by me. As shown in trans-sections 
Pl.y fig. Sf) the vagina, except its beginning portion, lies with 
in the internal layer of longitudinal muscles. • In this respect 
Tetracofrylus coryphicephalus agrees with the ffroteocephalidae . 


The ovary (PI. ^ fig.?/) is posterior and bilobed. fhe 
mid-piece connecting the lobes is narrow and thin, '^he lobes are 
long and somewhat broad. From the dorsal surface and at the distal 
region of each lobe rounded prominences arise and extend toward the 
dorsal surface of the proglottid. Many of these when observed in 
£oto preprations do not seem to have any connection with the ovary 
but when studied in trans-sections they are seen to be a part of it. 

Montioelli's drawing which is repfaCduced ( ?1.^ f ig.#^) 
does not truly represent this- appearance. 

These dorsal prominences and the greater part of the 
ovarian lobes (PI. fig. ) from which they arise lieK outside of 
the internal layer of longitudinal muscle;... The midpiece and apart 
of the lobes of the ovary lies within the muscular sheathe. 

JJcnticelli (1891) failed to point out this condition. 
Here is a relation the like of which I have been unable 


to find elsewhere in all the cestodes. So far as I know the ovary 
in other eestodes always lies in the medullay layer. 

At the posterior margin 01 the ovarian mid-piece is 
situated the oocapt. This organ was called the"siintere ovarieo" 
by Monticelli. The oocapt leads i£to the oviduct whcih after mak- 
ing several coils received the vagina. Then as a fertilization duct 
the oviduct continues until it reaches the o6type which is surrounded 
by many single shell glands. Just before entering the ootype the 
oviduct receives the unoaried vitelline duct. The ootype dis- 
charges into the uterine passage which passes ventral to the ovary 


and then discharges into the utjrus. It will be seen fron a 
comparison of the above description and Monticelli's drawing of 
those organs (PI. fig. ) with drawings and description of similar 
organs in the Pre teo c ephalidae that there is marked agreement be- 
tween T etracotylus and the Prof eoce p halidae in these relations. 
However this is no more than can be reasonably expected when it is 
remembered that practicably the same plan for these organs is 
followed thruout the whole order of Tetraphyllidea to which 
Tetracotylus and P rotoocephalus belong. 

Monticelli (1891) takes ocassion to point out what seems 
to be misinterpretation d>n the part of Von L ins tow (1891) in 
describing these organs in Taenia longicollis . From my own 
observations I agree with Monticelli in considering that V. Linstow 
has mistaken the oocapt for the ootype. A farther discussion of 
this point will be found in the description fo Proteocephalus 
lon?icollis . p. 

The vitellaria, which, in the Prot eocephalidae are lateral, 
follicular mas; es, the individual follicles of which are closely 
grouped about a central conducting tubule, ar^e in Tctraootylns 
coryphiceohalus (PI. a/ fig. ) widely extended lateral follicular 
masses which lie in a single ventral layer. In Proteoceohalids 
the vitellaria are within the longitudinal muscle layer while in 
— • coryphicephalus they lies outs| de f those muscles. She 

vitellaria are not only ventral but at the margin of the proglottid 
up toward the dorsal surface hence hwne viev/ed in a toto preparation 
the vitellaria at the margins seem greatly oompacted. 


A drawing ( PI. y fig. tff ) of a trans-section of the proglotti&s shows 
the method of arrangement o_ the vitellaria. The lateral long- 
itudinal vitelline ducts have not been seen on account of the 
poor histo : ogical condition of the material. The paired vitelline 
ducts however, may he seen (PI. fig. ) to arise in the lateral 
fields and to oass toward the middle of the proglottid outside 
of the longitudinal muscle layer. After running for some distance 
in this relation the ducts turn dorsally and enter 7/ithin this muscle 
layer and then they pass on to the middle of the proglottid where 
the paired vitelline duct which disch rges into the ootype. In all 
the specios of Proteocephalidae shown at present the vitelline 
ducts are always within the longitudinal muscle layer. Honticelli 
did not not'^ the position of the vitelline duets in coryphicephala. 

The uterus (PI. y fig.?/ ) in ripe proglot'ids is made 
up of about 18-30 slender ut pocketings, which extend well toward 
the margins of the proglottids but not as far lateral as in certain 

species of Proteocephalus such as ?. f allax , P. dub i us P. 
aiablopciitis ,etc. 


Monticelli (1891) states that the uterus when filled with eggs causes 
the ventral surface of the proglottid to belly out. Unfortunately 
this drawing showing that condition has not been repcf?duced. The 
uterus is ventral and it too lies outside of the internal layer of 
longitudinal muscles. This condition is well shown in the drawing 

(Pl.^ fig.frf ). 

I have not observed any uterine pores. Monticelli^ 
says that they do not exist but that when the uterus is full the 
ventral body wall ruptures, thus discharging the eggs. A careful 
study of good material would probably show the presence of one of 
more uterine | ores such as was found by Kraemer (1892), Schneider 
(1905) and by myself in a large number of Proteocephalids. 


EGGS.- Monticelli gives no measurements of the uerine 

eggs. He states that the membranes are quite delicate and that the 

uterine eggs are ve y small. In one of Monticelli' s slides one 

proglottidjj was broken open and some of the eggs were visible. The 
t tn b r if o s 

spheroidal ^measured about 0.015 to 0.017 ran. in diameter. The 
membranes had been rendered so transparent, by the balsam 7 ,hat they 
could not be measured. 

RELATIONSHIPS . - In the above description and in the 
drawings mentioned I have shown that Tetra co tylus coryphicep: ala 
differs from the Proteocephalidae and especial "y from the typical 


species of Proteocephalus in number, characters. These character 
concern the position of the genital • organs and their relation to the 
inner longitudinal muscle lawyer. In all the Proteooe anal i dag- 


which have "been sufficiently invest igated to determine these points 
the entire genital apparatus with the exception of a portion of the 
cirrus-pouch and vagina lies within the inner longitudinal muscle 
layer. In this family of the fol icles of the vitellaria are closely 
packed about a central duct. In Tetracotylus the vitellaria are in 
two broad lateral fields; the follicles of which are scattered and 
in a single layer. The vitelline ducts for some distance are outsid:; 
the inner muscles layer. The tests and uterus are entirely outside 
this muscle layer and the ovary is partly outside of it. As in the 
Proteocephalidae the vagina^vas deferens, ana part of ghe cirrus- 
pouch are within the inner muscle layer. On account of these marked 
difference between the Proteocephalidae and particularly the genus 
Proteocephalus . and the form which Monticelli named Tetracotylus 
o oryphic e ghal a % I propose to retain for the latter the generic name 
tohich Monticelli proposed for it, viz. Tetracotylus . The type of 
this genus is Tetracotylus coryphicophalus Lionticelli. The original 
ending of specific name is changed to agree with that of the generic 
name • 

The question of availability of the name Tetracotylus 
must be considered. 

Monticelli (1801) proposed Tetracotylu s as the name of 

a genus. He listed 2 species of fish cestodes which he believed 

belonged here and tho he did not designate a type, his genus was 


clearly based on T. Coryhice phala. 


Lbnnberg (l8 : J4j apparently ignored the reasons which Mon- 
ticelli gave for founding the genus and proposed for this group, 


the name Ichthyotaenia . Ichthyotaenia f ilicollis 1 was given as the 
type .and in the list of species he included I. ooryphioephala 
(Monticelli). RiggenbacH (1896) adopted Lonnberg's generic name 

"8 r a w n 

tho he knew of Monticelli's earlier name for the group. Bfotb 
(1894-1900:1680-81) stated, and I believe correctly, that the genus 
Tetracot;/lus Monticelli (1891) was based upon the species ttenia 
coryp hicephal* . This was equivalent to the declaration oi* a type 

Prior, however, to the appearance of Monticelli ! s (1891) 
work ffeinland (1858) proposed the generic name Proteo ce phalus and 
he made Taenia ambigua Du jar din the type of the new genus. He 
included in the genus Taeni a gllicollis and T. dispar . The latter 
species has been shown to "belong to a different genus, ITemato taenia* 
My investigations cause me to "believe that Taenia ambigua and Taenia 
f ilicollis are identical and for this reason Taenia ambigu a is a 
synonym of the older name Taenia f ilicollis Rud. and hence should 
be asrgpooc d. 

We inland and Tetraoolytus Monticelli 
If now the types of ? roteocephalus A are sufficiently alike 

in structure to belong to the same genus then Tetracotylus Monticelli 

would necessarily be considered a synonym of Proteocephalus 

Weinland and should be suppressed. I have shown, however, that the 

type of Tetracotylus differs greatly from the typical species of 

Proteocephalus and in my estimation these difference are sufficiently 

great to justify the separation of the species into different genera 

and moreover I consider that the two genera are so dissimilar in 



their internal organisations that they do not belong in the same 

For these reasons I retain the generic name Tetrao o tylu s 
Monticelli which Luhe (1899) and 3 : 4kn-( 1 C J00 ) consider ha;, been 
invalidated by reason of the earlier name Tetracotyle Filippi (1894). 
However T etraootylus and Tetracotyl a not "being spelled the same can 
not "be considered as homonyms and hence Tetraootylus is valid, 

3r<vttn (1894-1900) states that the spelling Tetraootylu s 
has "been used for Tetracot-rle and hence that name could not "be used 
"because it would then be a synonym of Tetracotyle . I have "been unable 
to find such a use of Tetraootylus in the literature at my disposal. 
If it has been so used prior to the time when Monticelli used it to 
designate this cestode genus then it is invalidated; if not until 
lator^then it is valid and ahould be retained. 

I propose to redefine the diagnosis of the genus Tetraoot ylus 
thus: Head somewhat globose, without folds or lappets encircling 
he$d or suckers. Ho true rostellum. Spines? Fifth sucker? Suckers 
four prominent, round or oval, margins of same entire. Heck usually 
long. Genital pore marginal, irregularly alternating. Cirrus-pouch 
and vagina discharging into a comon genital sinus. Genital papilla? 
Testes numerous scattered irregularly thruout a single field which is 
dorsal and outside of inner nuscles sheathe. Cirrus unarmed, slender 
? Vac deferens and vagina ithin inner muscle sheathe. Vitellarii 
Lateral , ventral composed of scattered follicles in cortical layer. 
Jterus with aay lateral outpo©ket ings , ventral, and in cortical layer 


Organs of interovarial space as in Proteocephalidae » 

In this genus protoably "belong Tetraeotylus Die singii 
Llonticelli and T. Hacrocotylea and Taenia maVlopteruri , FritsabL. 
These three speeies are all parasitic in the Siluridae t Perhaps 
here also "belongs Taenia osculata Goeae. 

I have perferred, however, for the present to consider 
this A as a species inquirenda i n the genus Proteocephalus in which 
it has long "been consider to "belong. I think it not unlikely that 
more careful research ilnto the morphology of Taenia o sculata w ill 
shov,- that it belongs to the genus Tetracotylu;-;,, 


TETRACOTYLUS DIESI1IGII Moiitioelli , sp. inf. 
(PI. 7 Pig. > 

1891: Tetracotylus Diesingl| Uonticelli , 1891. 

1896: Ichthyotacsia Diesingii Riggenbach, 1896:267. 

This species was very inadequately described by Llonticelli 
(1891) whose material was too limited and to poor to permit a 
careful study. Riggenbach (1896:267) listed it as one of the 
species of Ichthyotaenia . 

Monticellis ojigmnal description is here quoted: 

"TAENIA DIESIHGII.- Capo subgloboso anteriormente rotendato 
appena distinto dal collo. Le quattro ventose granc.i, pro eminent i 
e molto muscolari. Collo assai lungo : prime proglottidi brevissime 
appiattite, proglottidi mediane alquanto rigonfie all'aspetto 
carcinif ormi , posteriori rettangolari , ultimo subquadrate. Apeture 
genitali irregolarmente alternanti. L'apparato genitale, da quanto 
mi e riuscito veclere, si assomiglia a quello dalla T. coryphi c ephala . 

Habitat : intestino del 3ilurus dargado . " 

Thanks to Prof. H. B. 7/ard I have been able to examine 
Monticelli's type slides of this species and I can add the follow- 
ing data to the description by Monticelli. 

EXTERNAL CHARACTERS: The head is about 0.30 mm. in breadth. 
The suckers are about 0.17 mm. in their max dimension. The sucker 
opening is about 0.08 mm long. The musculature is thick. 

The neck is broken into several pieces. I judge that it is 
about 3-5 mm long. 


The young proglottids are about 0.0b mm long by 0.34 mm "broad. 
Fully mature proglottids are about 0.80 mm broad by 0.45-0.50 mm. 

The segmentation is not distinct. The proglottids are 
attached "by thier rail width. 

GENITAL ORGANS.- The genital pore is marginal situated 
within the first one-third or one-fourth of the proglottid. It 
alternates irregularly. 

Male organ s.- The cirrus-pouch is about 0.265 mm. long 
"by 0.105 ram broad. The ductus ejaculatorius forms numerous coils 
within the cirrus-pouch. The cirrus is not very muscular and is 
slender . It was not seen protruded. The testes are numerous 
probably about 100. They were situated in a single field. The 
coils of • as deferens could not be seen. 

Female organ s.- The vitellaria are lateral. The follicles 
seem to be scattered much as in T. Cor y o h i c e oh alu s . The ovary is 
posterior but its true character could not be termined. The vagina 
is anterior or posterior (?) to the cirrus-pouch. The uterus and 
eggs could not be observed in this material. 

RELATIONSHIPS.- From my examination of Monticelli's ma- 
terial I am satisfined that this species belongs in the genus 
Tetracotylus. it is closely related to T. c o r y? h ice ohalus but seem? 
tu differ, from that species. 7/ith the material at hand many 
points must remain in doubt. 


T etracotylus nacrocot ylea Monti cell i , sp. ino* 
(PI. Y fig. 87 ) 

1891: Tetracotylus rnacrocotylea Monticelli 1391: 
1896: I chthyotaenia rnacrocotyl e a Riggenbach 1896:267. 

This species was described by Mont' cell! (1391) Lack of 
sufficient good material did not permit a careful study of it. 
Rig~enbach (1896:267) listed this form in the genus Ichthyo taenia . 

Monticelli 's original description is here quoted: 

" Taenia rnacrocotylea .- Quest a Taenia, che indie o cosi dalla 
su prominenti ventose, ha molte rassomiglianze con le due seguenti, 
ma da entramoe si distingue per la caratteristica forma del capo 
che e nettamente distinto dal collo e per il collo piu breve. Le 
aperture genitali sono marginal! irregolarmente alternant i. Lo 
stato dell 1 individuo che posseggo non mi ha permesso uno studio 
piu minuzioso di questa Taenia. 

HABITAT: intestino del Silurus megace phalus . 

Prof. H. B. ".Yard very kindly secured Monticelli 's type 
slides of this species for my examination. As a result of this 
examination I can add the following data to the firinginal des- 

EXTER1TAL CHARACTERS. - The head is slightly crushed. In 
this condition it measures 1.10 mm broad a figure which probably 
too great by 0.10-0.20 mm. Its length is about 0.90 mm. 

The suckers are prominent and heavily muscled. They measure 
from 0.45 to 0.5U mm. in diarieter. 


The sucker cavity could not "be measured. 

The neck is about 0.50 mm broad just back of the head. 
Its length could not be ascertained for it was broken off. 

The mature proglottids are about 0,75 mm. square. Ho. 
young or ripe proglottids were seen. The segmentation is distinct. 

GENITAL 0RGAII3.- The genital pore is marginal, irregularly 
alternating and situated in the first one fourth of the proglottid. 
It is rendered prominent Iby a small papilla upon which it is 
placed. 2 he length of the cirrus-pouch and the relations of the 
latter with the vagina could not be ascertained, ^he testes are 
numerous, ab^ut 100 or more, and they are situated in a single field. 

Female organs. - The vitellaria are laterial, folliculor, 
and scattered much as they are in T_. c o r yph i c e ph alu s . The ovary 
is bilobed and posterior is but the matorial in two poor a condition 
to _ ermita study of it. The uterus could not be seen. 

RBLATI01ISHI2S.- This species, I am satisfied belongs 
in the genus Tetracoty'us. It is differentiated from the other 
species in the genus by its large head and suckers, its prominent 
genital pore and its evident begmentation. Unfortunately the 
mat rial is too poor to allow a careful study, which would make 
clear its exact relations to the other species of the genus. 


(PI. 9 lig§/?£ )/77j 

1886: Taenia malopt eruri Fritsch, 1386:103-8. 
18yi: T ctraco'cyl us malopteruri Llonticelli, 18y±; 
189b: lohthyoTa enia malopteruri Riggenbach, 1896:ii66. 

SPECIFIC DIAGNOSIS.- Characters of genus. Length of 
cestode ? Breadth up to 1.5 mm. Scolex fairly large, 0.45 mm. in 
"breadth, quadrangular , beaing four prominent suckers. Rostellum (?) 
hemisphaerical armed with many small hooks. Suckers large, heavily 
muscled^ protuberant f 0.25 mm. in diameter. Neck short, nearly broad 
as head. Mature proglottids nearly quadrate, Ripe proglottids 
broader than long, 1.46 mm. broad by 1.1 mm. long. 

GEHITAL ORG^SS.- As in genu . Genital pore marginal 
irregularly alternating, near middle of proglottid length. Pore 
large-, marked by thickening of margin of same. 

Male organs .- Cirrus when protruded, short and filiform. 
Cirrus-pouch short and relatively small, extending about 1/5 
across proglottid breadth, llo coils of ductus ejaculatorius in 
pouch. Vas deferens a mass of coils extending to middle of 
proglottid. Testes filling up field between vitellaria^about 120 
in number. 

Female organs .- Vagina posterior to cirrus-pouch. Ho 
coils of vagina anterior to ovary. Ovary a bilobed, winglike 
structure. Vitellaria f oil icular^ in lateral fields. Uterus a 
median tube with 18-20 lateral pouches. 


EGGS.- Not described, 

us (?) 
HABITAT.- In intestine Malopterui^ electricus A Egypt. 

HISTORICAL SUMMARY.- Fritsoh described and figured 
this form from Halopterurus electricu^ in 1886 as Taenia malopteruri. 
Mont ice Hi (1891) made this a species in his genus Tetracotylus . 
He did not describe it. Higgenbach (1896) considered this a species 
of Ichthyotaenia . 

LISCUSSIOIT.- The description given by Fritsch is very 
incomplete. However, he figured the head and a nearly rJpe proglottid 
givang the magnification of the same. I have used such data as 
could be secured from his description and have supplied the measure- 
ments needed from his drawings. Such measurement^ can not be very 
accurate yet they will probably be correct with in the limits of 
variation in the species. 

His diagnosis reads: "Taenia malopterurj » Caput q uid- 
rangulare, acetabulis ouattuor, angulariter posit is robustis. 
Rostellum hemisphaeEicum, spinis obtusis Teytuberoulis minimus 
ornat^um. Collura mediocre. Aperturae genitalium marginales leviter 
circumvallatae. Corpus sulco profundo per medium impressum. Articuli 
adulti dimido i'ere longiores quam lati, proglottideshreves, contract i 
Habitat in intestino Malopteruri. ■ 

BITERUAI CHARACTERS.- The length is not given. The 


breadth was up to about 1.5 mm. The seolox is of fair size 0.45 mm. 


in diameter 


It is quadrangular and "bears four prominent suckers and an 


:.rmed rostellum^ • The rostellum is hemisphaerical , armed with 
many small hooks. The suckers are large, 0.25 mm. in diameter. 
They are heavily muscled and are protuberant from the angles of the 

The neck is short and is nearly as "broad as the head. The 
proportions of first proglottids are not given. Mature proglottid s 
are nearly quadrate and ripe proglottids are somewhat "broader than 
long. Ripe proglottids measure about 1.46 mm. "broad "by 1.1 mm. 
long . r j 

GENITAL 0RGA1TS.- The sexual organs^are arranged according 

to type of genus. The genital pore is situated near the middle 

of the margin*- alternates irregularly from side to side • 
It is of large size and is marked by a considerable thickening of 
the margin. / , 

Male o rgans .- The cirrus^ is sometimes protruded and is 
then short and filiform. The cirrus-pouch is relatively small 
and short. Its length goes into the proglottid width about five 
times. There are no coils of ductus e jaculatorius v/ithin the 
cirrus-pouch. Outside the cirrus-pouch the coils of the vas 
deferens form a knot extending to the middle of the proglottid. 
The testes are irregularly scattered between the vitellaria and 
the ovary. Fritsch figures about 120 of them, H is drawing which is 
reproduced here (PI. / fig. /77 ) does not show them in the mid- 


Fenale organs . - The vagina^ opens posterior to tlio 
cirrus-pouch. Whether it ever opens anterior is not stated. 
There are no coils of the vagina anterior to the ovary. The ovary- 
is bilobed somewhat winglike and posterior . The lobes are ap- 
parently follicular at the outer ends. The vitellaria are long, 
follicular masses .in each lateral field. The uterus is a median 
tube from which arise many side-pockets. At least 18-20 on either 
side are shown in Fritsch's drawing. 

EGGS.- Hot described by Fritsch. 

P0SI3JI0H IB" CLASSIFICATION.- The position of this 

species is not readily determined from the data at hand. Fritsch' s 

description leaves much to be desired and his figures are not 

sufficiently definite to render a.positive determination possible. 

Nevertheless certain features of the anatomy of the worm are 

so well delineated that some definite conclusions can be reached. 

The rounded summit coverd with numerous spines and the very 

prominent suckers which seem to be set on the surface of the head 

are not" typical of Proteoce ohalus . Species of that genus lack 

spines and their suckers are never as prominent as in this species. 


The head agrees very well with that of Tetracylu s coryphicephalus 
Monti colli in regard to the suckers. The latter species has no 

In the genital organs there seems in the main to be a 
good agreement with the genus Proteocephalu s. Nevertheless careful 
comparison of Fritsch's drawing of the proglottid (PI. ? fig./77) 
with ray figure of Tetracotylu s (Pl.f fig. fl ) shows a certain 


resemblance which cannot be overlooked. In "both species there is 

the more or lees indefinite ovary m j? de up of follicles at the 

seem to be smaller than in P, coryphicephalus 
outer regions. These follicles in T. malopteruri^ ,The vi tell aria 

apparently are not as extensive in the former as in the latter yet 
from those of 

they are different A Proteocephalus . The lips of the 

genital sinus are much more prominent in this species than in 

Proteoceohalus, This character was not mentioned by Monticelli 

as being present in Tetracotylus nor have I noted it in his specimen;;* 

The species does not belong; to the genus Proteocephalus Weinland. 

I assign it with some reservation to the genus Tetracotylus 

Monticelli and the species is then to be shown as Tetracotylus 

malopteruri (Fritsch). 

The species of cestodes from the Siluridae merit much more 

carei'ul study by modern methods. In a study of the literature of 

this subject I noted the species ^tracggipos ciliotheca .Hfedl and 

UarsypocephaluB rectangulu s Vedl reported from Heterobranchus 

anguillaris b- "edl 1861 . The former on account of its ventral 


genital pore and ciliated embryo and tv;o* bothria evidently belongs 


to the Psejid^phyllidea * . The latter^ may belong somewhere 

in the ProteceTDhalidae or to Tetracotylus . Hot sufficient data 
are given to warrant a positive determination. Wedl (1861) also 
reported Scolex s;mdontis fro m Syndontis schal and from Hete ro- 
branchus anguillaris . These specimens likewise cannot be definitely 



- -» - 

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1900, 9n the Structure of Two Pish Tapeworms from the Genus 

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Observationes de Entozois. Gryphiswaldi 1#25. 
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1^50-1o51» Syetema helminthura. Vol. I und II. Vindobonae. 
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Histoire naturelle des Helminthes ou Vers Intestinaux, 

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1#£6. Die Parasiten des Zitterweises. Sitzungzber. d. konigl. 
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1#95» Die Tanien der Amphibien. Zool. Jahrb. Anat., IX, 207- 
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Fuhrmann, 0. 

1903- L' evolution des Tenias et en particulier. de la larve de 

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1#72. Zur Kenntnis einiger wenig bekannter Binnenwurmer. Nach- 
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- * - 

Plate 1 • 


1. Proteocephalus macrocephalue, ripe proglottid, female organs. 

2. n n " w male organs. 

3. " " head, to to. 

4. " n ripe proglottid, trans-section. 

5. " " neck folds, after Linton. 

6. Proteocephalus pentastomue, head and neck, n Kaptocz. 

7. " " middle of worm " " 
g. 11 n posterior end n " 

9. Proteocephalus exiguus, entire worm, " Benedict. 

-J Q tt If lift till 

11. n B head, toto, " " 

12. " " head, frontal 

section w " 

13. " tt proglottid, " n 

14. n " mature proglottid, toto. 

15. " " head, toto. 

16. " " ripe proglottid, toto. 

17. "" w end proglottid, toto. 
1#. Proteocephalus pusillus, mature proglottid, toto. 

19. " " ripe proglottid, toto. 

20. " " mature proglottid, toto. 

21. ■ " head, toto. 

22. " ■ head, " 

23. n n head, " 

Plate 2. 
- * - 



P r»o t e o c e t)ha lu 

f iliCGllJ.8 

whole worm. 




mature proglottid, toto. 




head, toto. 




ripe proglottid, toto. 




head, toto. 




young proglottid, toto. 




mature proglottid, toto. 

31 » 

a ,b » c w 


outline of proglottids, different 



Pro t e o c epha lu s 

f allax, 

" n M ant. and post,, 

after Kraemer. 




n n ripe " it ii 




head, toto, w " 




mature proglottid, toto, w w 




« ti t» 




ripe w n 




head, toto, 




ti it 




ripe proglottid, reconstruction 





" " w organs. 

Figure • 

Plate 3. 
- * - 


~L I \J uDUvv|41tllUu 

T»T Tip TiTTUT 1 Ot. f, 1 . t.O+.O . 
i lUv Ir www x*~l , w\J uU« 



mo tune nroerlott i . toto. 




old proglottid, toto. 




head, toto. 




tt ti 




head, frontal section, after Benedict 


a & b " 

n tt 

head & part of etrobila, w Leidy. 



ripe proglottid, 

frontal section- after* Bensdic - 




head, w n 

51 • 



head. toto. 




head. toto. 



strobila. after Mflller 




head. " M 




II It It 

56 . 



Tiieoe of stro"hi 1 a . n ** 




head. toto. 




fifth sucker, trans—section. 




ripe proglottid, partial reconstruct. 




ripe proglottid, toto. 

61 . 



head, frontal section. 

XXV U>VX p X> X VSXX UQfX W V V w JU Uii • 




ripe proglottid, toto. 




head, toto. 




ripe proglottid, toto. 




" w partial reconstruction. 


ria t,e 



- •si- 


Proteocephalus dubius, 

head, toto. 




n n 




n n 




young proglottid, ioio. 




ripe proglottid, toto. 




mature progioivbid, t-olo. 




head, toto, after Zschokke. 




•> tt n it 




progio t uici , 


Proteocephalus ijjmlosus, 

ripe proglottid., uo»*o-» 




head, toto. 




ripe proglottid, after Kraemer. 




" w frontal section. 




w " trans-section. 


Proteocephalus longicollis, 

" " after von Linstow, 



cirrus-pouch w n tt 




ripe proglottid, 

J* — _ , _ _L _J ft ft ft 



Proteocephalus neglectus, 

ripe proglottid. 




cirrus-pouch and vagina, normal. 




w tt tt tt 

after Monticelli, 




head " 




n it n 



ripe proglottid, toto, 




w trans-section, 




head, after Monticelli. 
ripe proglottid, toto. 

P 1 n t p . 

Figure • 

92. Proteooephalus 


head, toto. 




n it 




cirrus-pouch and vagina, reconstruct. 




ripe proglottid, frontal section. 



tt it ti it 




head, trans-section. 




ti ti « 




w frontal " 




it tt tt 

101 . 



" trans- M 

102. Proteooephalus 


ripe proglottid, toto. 




head, toto. 




it it 




n it 




head and regions of strobila. 




proglottid, trans-section. 




head, frontal section. 




uterine pores unru cuoicuia. 

1 10. 



cirrus, vas deferens, and vagina, 




excretory ducts to exterior. 



head, toto. 




tt tt 

ria bo . 


— ™ 

1 14. 


f ilaroides, 

end organ or vestigial fifth 


sec uion. 




it « tt « tt 

tt n 

1 16. 



N tt It fl tt tt 





tt It ft ft tt tt 

n tt 




fieaci ox slquxu^ x»ooo» 




neaci 01 pierocoi ous^ iiOw« 




it tt n tt 




y*1 tip TTf ncl + + n / 1 +• /-\ -f- - 




Ilia.> pX UgXUliOXU) ovJuU* 




organs of inter-ovarial space 
s u mo x» x on • 

, re con- 



Tilria -r> t»o clot. t T rl trttn 



X HI ivj. J. f 

IXlck uUX O ^JXvj^XL/uuXUlj liU uUi 





X ClX I<~L 10^ 

UX ^ettXltJ \JX XIXOOX — U V CtX Xal OLJiAOtJ 

f L L UIIl L/U OU • 




cirruB-pouon ana. vagina > 

tt tt 




tt tt ti tt 

tt It 




lit; {ACL j TjtOLO. 




indentation at genital pore. 



132. Ophiotaenia calmettei, mature proglottida, toto. 

•j ^ ^ if it ti n ti 

134. " " head, toto. 

135* Ophiotaenia trimeresuri, cirrus and cirrus-pouch, from toto 


<J2£ # tt « It ft « n Mtlfl 

137* n " mature proglottid, toto. 

13#« Crepidobothrium gerrardii,head, toto, after Smith. 

1^9 # n w tin tin 

140. " " proglottid, toto. 

141. " " cirrus and vagina, after Monticelli. 

142. " " ripe proglottid, toto. 

143. " " mature " " 
1 Vi w « 11 11 ii n 

145. " " young " w after Smith. 

146. " ■ head, toto. 

147. " ■ ■ ■ 

143. " " mature proglottid, toto. 

Plate 5. 





proglottid, toto, 

after Parona. 




head, n 






n tt 






proglottids, n 




Ophiotaenia marenzelleri,head n 





mature proglottid, 






Crepidobothrium gerrardii, head, toto, 






tt n 












ripe proglottids 






head, toto, 














young proglottids, 







mature proglottid, 
















v. L ins tow. 











mature proglottid, 














nearly ripe proglottid 

n 11 





head, toto, 


Schwarz* . 





it tt 






ripe proglottid 



- * ■ 

j * 




proglottids, toto, after Klaptocz. 




ripe proglottid, " " 





head, " " 





« w tt 




lllcX-LUy utj I Hi X , 

n tt n 





ripe proglottid, w n 




Sal vo J. XII X j 

head, " " 





« tt it 




BgUHl , 

proglottid, " " 


1$1 . 



head, w " 



(?) simplicissimus, 

head & proglottids, Bn 



Choanoscolex abscisus, 

head toto," 





tt n tt 

v. Linstow. 


Proteoc ephalus 


n tt it 

tt « 




ripe proglottid, tt 

ti tt 


Proteocephalus fossatus, 

male organs, n 




head, toto," 





ripe proglottid, w B 





proglottid, " n 



Tetracotylus alternatim transverse lineata, head, toto, after Goeze. 



it « 

w regions of 

after Goeze. 



« n 

" posterior proglottids, 
toto, after Goeze 



osculatus, young, toto, after Wagoner from Carus. 




head, n " 

p£. / 




6 . 3 






1/ /S " /6 



2.2- 23 


PI 2. 



PI 2- 

Hi 5V 





6 2~ 


6 b' 


PI 5 

a w 70 



7ZT *o 







^ *7 




n ^ 




Pi 4 




Pi 7 

13 b' 






Wl /J7 


Ms- ftf 


Pi 7 

Pi 8 



PI 8 


17* , 7 f 


/*/ 195 m 


/?/ l<jz~ IH2> im 

w\ ** x «\ 


George Roger LaRue. 
- * - 

1g£>2: Born at Paulina, Iowa. 
1 ££7-1 £97: Attended public schools at Paullina, Iowa. 
1&9&-1900: Student at Crete Academy, Crete, Nebraska. 
1900-1901 & 

1903-1907: student at Doane College, Crete, Nebraska. 

1904-1907: Student Assistant in Biology, Doane College. 

1907: B. S. Doane College, with honorB of class. 

1907-1909: Graduate Student in Zoology, University of Nebraska, 
Lincoln, Nebraska. 
Technician in College of Medicine, University of Nebraska 

190&: Sigma Xi, Nebraska Chapter. 

1909: A. M. in Zoology, University of Nebraska. 

Thesis: "On the Morphology and Development of a new Ces- 
tode of the Genus Proteocephalus We inland", published 
Trans. A. M. S. 1909, pp. 17-46, 4 plates, 49 figures, 

1909-191 1: Graduate Student in Zoology, University of Illinois, 
Urbana, Illinois. 
Research Assistant in Zoology. 

1910: Elected to membership in the Gamma Alpha Graduate Scient- 
ific Fraternity, Illinois Chapter. 

Member of Illinois Academy of Science. 

Assistant, Biological Summer School of University of 
Illinois at Havana, Illinois. 

1 , - '